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DISTANCE LEARNING CENTRE

AHMADU BELLO UNIVERSITY


ZARIA, NIGERIA

COURSE MATERIAL

FOR

Course Code & Title: HPHY 225 (HUMAN PHYSIOLOGY I)

Programme Title: Bachelor in Nursing Sciences (B.N.Sc)

1
ACKNOWLEDGEMENT

We acknowledge the use of the Courseware of the National Open University of


Nigeria (NOUN) as the primary resource. Internal reviewers in the Ahmadu Bello
University who extensively reviewed and enhanced the material have been duly
listed as members of the Courseware development team.

2
COPYRIGHT PAGE

© 201 8 Distance Learning Centre, Ahmadu Bello University, ABU Zaria, Nigeria

All rights reserved. No part of this publication may be reproduced in any form or by any means,
electronic, mechanical, photocopying, recording or otherwise without the prior permission of the
Director, Distance Learning Centre, Ahmadu Bello University, ABU Zaria, Nigeria.

First published 2018 in Nigeria.

ISBN:

Published and printed in Nigeria by:

Ahmadu Bello University Press Ltd.

Ahmadu Bello University,

Zaria, Nigeria.

Tel: +234

E-mail:

3
ABU DLC COURSE WRITERS/DEVELOPMENT TEAM

Mohammed Hadiza Sani DLC Subject Matter Reviewer


Ene Adakole DLC Language Reviewer
Dr. Fatima Kabir
Nasiru Tanko DLC Instructional Designers/Graphics
Prof. A. Z. Hassan DLC Editor

4
CONTENTS

Title Page…………………………………………………………….………1
Acknowledgement Page………………………………………….… ……...2
Copyright Page………………………………………………………..…….3
Course Writers/Development Team……………………………………….4
Table of Content………………………………………………….................5
i. Course Information 7
ii. Course Introduction and Description 7
iii. Course Prerequisites 8
iv. Course Learning Resources 8
v. Course Objectives and Outcomes 8
vi. Activities to Meet Course Objectives 9
vii. Time (To complete Syllabus/Course) 10
viii. Grading Criteria and Scale 11
ix. OER Resources 12
x. ABU DLC Academic Calendar 15
xi. Course Structure and Outline 16
xii. STUDY MODULES 20
Module 1: General Physiology ………………………………………..……20
Study Session 1: Cell ………………………………………………..20
Study Session 2: Transport across Cell membrane ………………….35
Study Session 3: Biologically Important Molecules and Their
Functions ………………………………………….49
Study Session 4: Homeostasis……………………………………….55

Module 2: Nerve and Muscle Physiology, Blood


Physiology (I) - Haemodynamics and Immunology…………..70
Study Session 1: Nerve and Muscle Physiology ……………………70
Study Session 2: Blood and Body Fluids ……………………………89
Study Session 3: Haemostasis …………………………………….....111

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Study Session 4: Immune System …………………………………..119

Module 3: Neurophysiology I………………………………………………128


Study Session 1: Physiology of the Central Nervous System:
Brain ………………………………………………128
Study Session 2: The Spinal Cord, Cranial and Spinal Nerves …….147
Study Session 3: Neurons and Neuronal Transmission …………….165
Study Session 4 & 5: The Autonomic Nervous System ……………192

xiii. GLOSSARY……………………………………………………..209

6
COURSE STUDY GUIDE
I. COURSE INFORMATION
Course Code: HPHY 225
Course Title: Human Physiology I
Credit Units: 2 Credit Units
Year of Study: Two
Semester: First

II. COURSE INTRODUCTION AND DESCRIPTION


Introduction:
Let us start by defining physiology as the scientific study of how the body works
under normal conditions or in a state of good health. It describes how cells
operate, how they combine their functions in specific organs, and how these
organs work together to maintain a stable environment inside the body. We can
rightly say that physiology is the functional basis of the health sciences, because
most disease states are the result of disturbances of physiological processes.

Description:
A basic knowledge of physiology is essential for all students such as you whose
professional careers will involve aspects of health and patient care. Physiology is
also one of the key subjects in biomedical sciences and continues to be at the
forefront of biomedical research. Human Physiology (I) is the first of two courses
that runs in the second year of your programme. This course covers cell
physiology, maintenance of homeostasis, muscle function, body fluids and the
immune system.

7
III. COURSE PREREQUISITES
The prerequisites include the following:- Attendance of 95% of all interactive
sessions, submission of all assignments to meet deadlines; participation in all
CMA, attendance of all laboratory sessions with evidence as provided in the log
book, submission of reports from all laboratory practical sessions and attendance
of the final course examination. You are also expected to have:
1. Satisfactory level of English proficiency
2. Basic Computer Operations proficiency
3. Online interaction proficiency
4. Web 2.0 and Social media interactive skills

IV. COURSE LEARNING RESOURCES

(i) Fox, S.I. (2012). Human Physiology. 12th edition, Mc Graw Hill, NewYork.

(ii) Ganong,WF. (2010). Review of Medical Physiology. 23rd edition,

Mc Graw Hill, NewYork.


(iii) Guyton,A.C., Hall, J.E. (2001). Textbook of Medical Physiology.

Harcourt International Edition,10th edition, W.B.


Saunders,Philadelphia.
(iv) Oyebola, D,O. (2002). Essential Physiology, Vol 1, NihortPress.

V. COURSE OBJECTIVES
This course is aimed at ensuring that you can:
i. Discuss the context of the cell as the functional unit of the body.
ii. Apply the understanding of the mechanisms of dynamics of
body fluids, homeostasis and the immune process in

8
understanding changes and the control of the physiological body
process of patients.

VI. ACTIVITIES TO MEET COURSE OBJECTIVES


Read and understand the context of this course by reading through this course
guide and paying attention to details. You must know the requirements before
you will do well.

Develop a study plan for yourself. Follow instructions about registration and
master expectations in terms of reading, participation in discussion forum, end of
unit and module assignments, laboratory practical and other directives given by the
course coordinator, facilitators and tutors.

Read your course texts and other reference textbooks.


Listen to audio files, watch the video clips and consult websites when given.
Participate actively in online discussion forum and make sure you are in touch
with your study group and your course coordinator.
Submit your assignments as at when due.
Work ahead of the interactive sessions.
Work through your assignments when returned to you and do not wait until when
examination is approaching before resolving any challenge you have with any unit
or any topic.
Keep in touch with your study centre, the DLC ABU, websites as information
will be provided continuously on these sites. Be optimistic about doing well.

Specifically, this course shall comprise of the following activities:


1. Studying courseware
9
2. Listening to course audios
3. Watching relevant course videos
4. Course assignments (individual and group)
5. Forum discussion participation
6. Tutorials (optional)
7. Semester examinations (CBT and essay based).

VII. TIME (TO COMPLETE SYLABUS/COURSE)


The course will be delivered adopting the blended learning mode, 70% of online
but interactive sessions and 30% of face-to-face during laboratory sessions. You
are expected to register for this course online before you can have access to all
the materials and have access to the class sessions online. You will have hard
and soft copies of course materials, you will also have online interactive
sessions, face-to-face sessions with instructors during practical sessions in the
laboratory. The interactive online activities will be available to you on the course
link on the Website of DLC ABU. There are activities and assignments online
for every unit every week. It is important that you visit the course sites weekly
and do all assignments to meet deadlines and to contribute to the topical issues
that would be raised for everyone’s contribution.

You will be expected to read every module along with all assigned readings to
prepare you to have meaningful contributions to all sessions and to complete all
activities. You are expected to commit a minimum of 2 hours daily for this
course.

10
VIII. GRADING CRITERIA AND SCALE
Grading Criteria
A. Formative assessment
Grades will be based on the following:
Individual assignments/test (CA 1,2 etc) 20
Group assignments (GCA 1, 2 etc) 10
Discussions/Quizzes/Out of class engagements etc 10

B. Summative assessment
CBT based 30
Essay based 30
TOTAL 100%

C. Grading Scale:
A = 70-100
B = 60 – 69
C = 50 - 59
D = 45-49
F = 0-44

D. Feedback
Courseware based:
1. In-text questions and answers (answers preceding references)
2. Self-assessment questions and answers (answers preceding references)

Tutor based:
1. Discussion Forum tutor input
11
2. Graded Continuous assessments

Student based:
1. Online programme assessment (administration, learning resource,
deployment, and assessment).

IX. Open Education Resources


OSS Watch provides tips for selecting open source, or for procuring free or open
software.
SchoolForge and SourceForge are good places to find, create, and publish open
software. SourceForge, for one, has millions of downloads each day.
Open Source Education Foundation and Open Source Initiative, and other
organisation like these, help disseminate knowledge.
Creative Commons has a number of open projects from Khan
Academy to Curriki where teachers and parents can find educational materials for
children or learn about Creative Commons licenses. Also, they recently launched
the School of Open that offers courses on the meaning, application, and impact of
"openness."
Numerous open or open educational resource databases and search engines
exist. Some examples include:
i. OEDb: over 10,000 free courses from universities as well as reviews of
colleges and rankings of college degree programmes
ii. Open Tapestry: over 100,000 open licensed online learning resources for an
academic and general audience

12
iii. OER Commons: over 40,000 open educational resources from elementary
school through to higher education; many of the elementary, middle, and
high school resources are aligned to the Common Core State Standards
iv. Open Content: a blog, definition, and game of open source as well as a
friendly search engine for open educational resources from MIT, Stanford,
and other universities with subject and description listings
v. Academic Earth: over 1,500 video lectures from MIT, Stanford, Berkeley,
Harvard, Princeton, and Yale
vi. JISC: Joint Information Systems Committee works on behalf of UK higher
education and is involved in many open resources and open projects
including digitising British newspapers from 1620-1900!
Other sources for open education resources
Universities
i. The University of Cambridge's guide on Open Educational Resources for
Teacher Education (ORBIT)
ii. OpenLearn from Open University in the UK
Global
i. Unesco's searchable open database is a portal to worldwide courses and
research initiatives
ii. African Virtual University (http://oer.avu.org/) has numerous modules on
subjects in English, French, and Portuguese
iii. https://code.google.com/p/course-builder/ is Google's open source software
that is designed to let anyone create online education courses
i. Global Voices (http://globalvoicesonline.org/) is an international community
of bloggers who report on blogs and citizen media from around the world,
including on open source and open educational resources

13
Individuals (which include OERs)
i. Librarian Chick: everything from books to quizzes and videos here, includes
directories on open source and open educational resources
ii. K-12 Tech Tools: OERs, from art to special education
iii. Web 2.0: Cool Tools for Schools: audio and video tools
iv. Web 2.0 Guru: animation and various collections of free open source
software
v. Livebinders: search, create, or organise digital information binders by age,
grade, or subject (why re-invent the wheel?)
Legal help
i. New Media Rights is trying to help digital creators use public domain or
open materials legally. They have guides on how to use free and open
software materials in various fields.

14
X. ABU DLC ACADEMIC CALENDAR/PLANNER

PERIOD
Semester Semester 1 Semester 2 Semester 3
Activity JAN FEB MAR APR MAY JUN JUL AUG SEPT OCT NOV DEC
Registration
Resumption
Late Registn.
Facilitation
Revision/
Consolidation
Semester
Examination

N.B: - All Sessions commence in January


- 1 Week break between Semesters and 6 Weeks vocation at end of session.
- Semester 3 is OPTIONAL (Fast-tracking, making up carry-overs & deferments)

15
XI. COURSE STRUCTURE AND OUTLINE

Course Structure
WEEK/DAYS MODULE STUDY SESSION ACTIVITY

1 Study Session 1: 1. Read Courseware for Study Session 1.


Title: The Cell 2. View the Video(s) on Study Session 1
p. 20 3. Listen to the Audio on Study Session 1
4. View any other Video/U-
tube(https://www.youtube.com/watch?v=gG7uCskUOrA
5. https://www.youtube.com/watch?v=JulrOABEKIY )

2 Study Session 2 1. Read Courseware for Study Session 2.


Title: Transport 2. View the Video(s) on Study Session 2
across Cell 3. Listen to the Audio on Study Session 2
membrane 4. View any other Video/U-tube(https://youtu.be/J5pWH1r3pgU )
p.35

3 Study Session 3 1. Read Courseware for Study Session 3.


Title: Biologically 2. View the Video(s) on Study Session 3
important molecules 3. Listen to the Audio on Study Session 3
and their functions 4. View any other Video/U-tube(https://youtu.be/FgfknBZaVTI)
STUDY p.49

4 1. Read Courseware for Study Session 4.


MODULE 1 Study Session 4 2. View the Video(s) on Study Session 4
Title: Homeostasis 3. Listen to the Audio on Study Session 4
and Nerve and 4. View any other Video/U-tube(https://youtu.be/Iz0Q9nTZCw4
Muscle Physiology https://youtu.be/Jyh8HhK80X8?list=PLB73vUwd2zzYSiuGydp8G2fEEbjo7v
p.55 oDdhttps://youtu.be/0mhAN4-
8uWo?list=PLEsDFsi9ffTztzB5qDft5Ypn8RTJ5Ev8O)

5 1. Read Courseware for Study Session 1.


2. View the Video(s) on Study Session 1

16
Study Session1 3. Listen to the Audio on Study Session 1
Title: Body Fluids I
p.70

6 1. Read Courseware for Study Session 2


STUDY Study Session2 2. View the Video(s) on Study Session 2
Title: Body Fluids II 3. Listen to the Audio on Study Session 2
MODULE 2 p.89 4. View any other Video/U-tube(ihttps://youtu.be/tC2gMDfzYA8)

7 Study Session3 1. Read Courseware for Study Session 3.


Title: Haemostasis 2. View the Video(s) on Study Session 3
p.111 3. Listen to the Audio on Study Session 3

8 Study Session 4 1. Read Courseware for Study Session 4.


Title: Immune 2. View the Video(s) on Study Session 4
System. 3. Listen to the Audio on Study Session 4
p.119 4. View any other Video/U-
tube(https://youtu.be/Xc_Ljc5ycfMhttps://youtu.be/BFeJlxIldAI

9 Study Session 1 1. Read Courseware for Study Session 1.


Title: Physiology of 2. View the Video(s) on Study Session 1
the Central Nervous 3. Listen to the Audio on Study Session 1
System: Brain 4. View any other Video/U-
p.128 tube(https://www.youtube.com/watch?v=q8NtmDrb_qohttps://www.youtube.
com/watch?v=dHURMD4v8Kk)

10 Study Session 2 1. Read Courseware for Study Session 2.


Title: The Spinal 2. View the Video(s) on Study Session 2
Cord, Cranial and 3. Listen to the Audio on Study Session 2
Spinal Nerves 4. View any other Video/U-tube(https://www.youtube.com/watch?v=Sl5Z2-
STUDY p.147 STBRU
https://www.youtube.com/watch?v=t3i5sAVR3Wg
MODULE 3 https://www.youtube.com/watch?v=eYVzdDWSQdc
https://www.youtube.com/watch?v=6ENCJkXJvio)

17
11 1. Read Courseware for Study Session 3.
Study Session3 2. View the Video(s) on Study Session 3
Title: Neurons and 3. Listen to the Audio on Study Session 3
Neuronal 4. View any other Video/U-
Transmission tube(https://www.youtube.com/watch?v=WhowH0kb7n0
p.165 https://www.youtube.com/watch?v=vyNkAuX29OUhttps://www.youtube.co
m/watch?v=cUGuWh2UeMk
https://www.youtube.com/watch?v=-9xBfvh7hgg
https://www.youtube.com/watch?v=gc3jJKxlHyw

12 Study Session 4 & 1. Read Courseware for Study Session 4.


5 2. View the Video(s) on Study Session 4
The Autonomic 3. Listen to the Audio on Study Session 4
Nervous System 4. View any other Video/U-
p.192 tube(https://www.youtube.com/watch?v=7dZHmKMLdC0
https://www.youtube.com/watch?v=TDgD2ZE7_S4

Week 13 ON-CAMPUS TUTORIALS/REVISION

Week 15 & 16 SEMESTER EXAMINATION

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XII. Course Outline
1.1 Module 1: General Physiology
Study Session 1: Cell
Study Session 2: Transport across Cell membrane
Study Session 3: Biologically important molecules and their functions
Study Session 4: Homeostasis

1.2 MODULE 2: Nerve and Muscle Physiology, Blood Physiology (I) -


Haemodynamics and Immunology
Study Session 1: Nerve and Muscle Physiology
Study Session 2: Blood and Body Fluids
Study Session 3: Haemostasis.
Study session 4: Immune System

1.3 MODULE 3: Neurophysiology I


Study Session 1: Physiology of the Central Nervous System: Brain
Study Session 2: The Spinal Cord, Cranial and Spinal Nerves
Study Session 3: Neurons and Neuronal Transmission
Study Session 4 & 5: The Autonomic Nervous System

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STUDY MODULES
MODULE 1: General Physiology
Contents:
Study Session 1: The Cell
Study Session 2: Transport across Cell membrane
Study Session 3: Biologically important molecules and their functions
Study Session 4: Homeostasis

STUDY SESSION 1
The Cell
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
1.1 Organisation of the cell
1.2 Cell Structure
1.3 Cell or Plasma Membrane
3.0 Conclusion/Summary
4.0 Self-Assessment Questions and Answers
5.0 Additional Activities (Videos, Animations & out of Class activities)
6.0 References/Further Readings

Introduction:
As we said earlier in the Introduction, physiology is the scientific study of how
the body works under normal conditions or in a state of good health. It describes
how cells operate, how they combine their functions in specific organs, and how
20
these organs work together to maintain a stable environment inside the body.
Physiology is the functional basis of the health sciences, because most disease states
are the result of disturbances of physiological processes. A basic knowledge of
physiology is therefore essential for you whose professional careers will involve
aspects of health and patient care. Physiology is also one of the key subjects in
biomedical science and continues to be at the forefront of biomedical research.
Human Physiology (I) is the first of two courses that runs in the second year of your
programme. This course covers the cell physiology, the maintenance of
homeostasis, muscle functioning and the immune processes.

The human body is made up of several cells that perform basic functions that
sustain life. Different types of cells aggregate to form organs that ultimately
perform different functions. While different organs perform different functions, the
body must function in harmony. This module covers cell functioning and body’s
methods of achieving harmony through homeostasis. The basic living unit of the
body is the cell. Each organ is an aggregate of many different cells held together by
intercellular supporting structures. Each type of cell is specially adapted to perform
one or a few particular functions. For instance, the red blood cells, numbering 25
trillion in each human being, transport oxygen from the lungs to the tissues.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Discuss how the cell performs the various functions.
2. Discuss how the plasma membrane performs its functions
3. Discuss how the body sustain homeostasis with contribution from the different
body systems
21
4. Explain the Organisation of the cell
5. Explain 3 different substances that make up the cell
6. Give detailed explanation of the Cell Structure
7. Draw a typical cell showing the organelles in the cytoplasm and the nucleus.
8. Explain the functions of the following concepts- (a) Nucleus (b) Cytoplasm (c)
Endoplasmic reticulum and ribosomes (d) Golgi apparatus (e) Mitochondria
(f) Centrosome (g) Lysosomes
9. Describe the Cell or Plasma Membrane

2.0 Main Content


2.1. Organisation of the Cell
The cell has two major parts namely the nucleus and the cytoplasm. The nucleus is
separated from the cytoplasm by a nuclear membrane, and the cytoplasm is
separated from the surrounding fluids by a cell membrane, also called the plasma
membrane. The different substances that make up the cell are collectively called
protoplasm. Protoplasm is composed mainly of five basic substances: water,
electrolytes, proteins, lipids, and carbohydrates.

Proteins
After water, the most abundant substances in most cells are proteins, which
normally constitute 10% to 20% of the cell mass. These can be divided into two
types: structural proteins and functional proteins.

Lipids
The biologically important lipids are the fatty acids, triglycerides, phospholipids and
sterols. Fatty acids can be saturated or unsaturated while phospholipids are found in
22
cell membranes where they act as a structural component. Fatty acids also serve as
an important source of energy in the body.
Carbohydrates
Carbohydrates are organic molecules made up of equal amounts carbon and water.
They perform both structural and functional roles. They also help in cell signalling.
ITQ 1: The cell has two major parts namely ……
ITA 1: The nucleus and the cytoplasm

2.2. Cell structure


The cell is not merely a bag of fluid, enzymes, and chemicals; it also contains
highly organised living structures, called intracellular organelles. The physical
nature of each organelle is as important as the cell’s chemical constituents for cell
function. For instance, without one of the organelles, the mitochondria, more than
95% of the cell’s energy realised from nutrients would cease immediately. The
most important organelles and other structures of the cell are shown in Figure 1.1.1

Figure 1.1.1: A typical


cell, showing the
organelles in the
cytoplasm and the
nucleus.

23
Nucleus
The nucleus, shown in Figure 1-2, is usually a spherical organelle, though its shape
may vary in some cells. It is surrounded by a membrane called nuclear membrane.
The nuclear membrane has double layer and the two layers are fused at some points
to produce nuclear pores which are thought to allow molecules pass between the
nucleus and cytoplasm. There is a smaller spherical structure within the nucleus,
the nucleolus. The fluid contained within the nucleus is called nucleoplasm to
differentiate it from the fluid in the rest of the cell which is referred to as
cytoplasm. The nucleus is best seen in a cell that has been stained because the
chromatin within the nucleus stains vividly. In the unstained state, chromatin is
colourless. Chromatin gives rise to the chromosomes when a cell divides.
Chromosomes are primarily composed of deoxyribonucleic acid (DNA). DNA is
the basic substance for inheritance. The second basic substance of inheritance is
ribonucleic acid (RNA) which is generally contained within the nucleolus.
During cell division, genetic information contained in DNA is transferred to RNA,
which carries the genetic information out of the nucleus into the cytoplasm where it
directs the formation of protein. Thus, the nucleus is the repository of genetic
information for our whole body.

24
Figure 1.1.2: Structure of the nucleus

Cytoplasm
This is the jelly-like fluid between the nuclear membrane and the cell
membrane inside which the cell organelles are suspended. The clear liquid portion
in which the particles are suspended is called the cytosol. Cytoplasm is mostly water
but it contains electrolyte and about 15% protein plus fat and carbohydrate. The
cytoplasm comprises about 80% of the total weight of the cell.

Endoplasmic Reticulum and Ribosomes


There are small cytoplasmic tubules collectively called the endoplasmic reticulum.
Some tubules of the endoplasmic reticulum have small, spherical structures called
ribosomes attached to their membranes. Where these are present, the reticulum is
called the granular or rough endoplasmic reticulum. Other tubules are free of
ribosomes. This part is called the granular, or smooth, endoplasmic reticulum, as
shown in Figure 1.3. The tubules of the granular, ribosome-containing endoplasmic
25
reticulum are involved in the vital processes of protein synthesis and secretion in
the cell. A molecule of RNA is formed from the DNA in the nucleus. This RNA is
known as messenger RNA (mRNA) because it carries the genetic message from
DNA in the nucleus and passes through the pores in the nuclear membrane to enter
the cytoplasm. The mRNA then becomes attached to ribosomes, where it directs the
formation of proteins. Once synthesised, protein enters the tubules of the
endoplasmic reticulum. After the protein accumulates in the tubules, parts of the
tubules break off to become spherical vesicles containing quantities of protein.
These vesicles then become part of the golgi apparatus and the protein is
eventually secreted from the cell.

The granular endoplasmic reticulum does seem to be involved in protein synthesis;


yet many hormones are found stored in these tubules. In the cells of glands that
secrete hormones, such as the thyroid gland, the granular endoplasmic reticular
tubules contain large quantities of the hormone. Smooth endoplasmic reticulum is
involved in the synthesis of lipid steroid hormones and, in liver cells, contains the
enzymes which catalyse glycogen breakdown.

Figure 1.1.3: Structure of


the endoplasmic reticulum

26
Golgi apparatus
The Golgi apparatus, also referred to as the Golgi complex or Golgi body, shown in
the Figure, appears as a collection of tubules and vesicles. Secretory granules are
formed in the Golgi apparatus. These granules are packages of highly concentrated
protein. Once protein has been formed by the ribosomes, it accumulates in the Golgi
apparatus where it is concentrated and may be modified and then packaged into
vesicles of secretory granules. These vesicles fuse with the membrane and then
open up to release the protein from the cell. Carbohydrate may be added to the
protein within the Golgi body to form glycoproteins. Mucus is also formed in this
area.

Figure 1.1.4: A typical Golgi apparatus and its relationship to the endoplasmic reticulum (ER)
and the nucleus.

27
Mitochondria
The mitochondrion, shown in the Figure, is called the “powerhouse” of the cell.
Without the mitochondria, cells would be unable to extract enough energy from the
nutrients, and essentially all cellular functions would cease. In the mitochondria, the
very important compound adenosine triphosphate (ATP) is formed. ATP is said to
be a high-energy phosphate compound because, when it splits off a phosphate
molecule to become adenosine diphosphate (ADP), energy is made available to the
cell. This energy is used for the various cellular processes, such as the contraction
of a muscle cell.

Mitochondria are found in varying numbers in all nucleated cells. They may be
distributed evenly throughout the cytoplasm or concentrated in areas of high
energy requirement; for example they lie between the fibrils of muscle fibres
where they produce energy for contraction. Each mitochondria is bounded by a
smooth outer membrane which is separated by a small space of about 8nm from a
folded inner membrane. These folds are called cristae and they are studded with
minute particles. The inner and outer membranes, the space between them, the
membrane bound particles and the inner matrix contains enzymes. All of the
enzymes which break down nutrient substances into carbon dioxide and water,
together with the enzymes which enable the transfer of released energy to stable
high-energy compounds are present within the mitochondrial structure. Virtually all
of the cell’s high-energy compounds are synthesised within the mitochondria.

Mitochondria can increase its own number by repeated self-replication. This occurs
when there is need for increased amounts of ATP in the cell. Mitochondria contain
28
deoxyribonucleic acid (DNA) similar to that found in the nucleus. The DNA of the
mitochondrion, like that in nucleus, controls its replication.

Figure 1.1.5: Structure of Mitochondria

Centrosome
The centrosome contains two centrioles. They lie close to the nucleus. At the
beginning of cell division, the two centrioles divide, thus forming four centrioles,
one pair goes to one end of the cell and the other pair to the opposite end. The
centrioles function to pull the chromosome pairs apart. In this way, one set of
chromosomes goes to one side of the cell and the other set to the other. When the
cell divides, each new cell has a complete set of chromosomes.

Lysosomes
Lysosomes are vesicular organelles that form by breaking off from the Golgi
apparatus and then dispersing throughout the cytoplasm. When a cell engulfs
bacteria, the bacteria come in contact with the lysosomal enzymes, which then

29
destroy them. When a cell dies, the lysosomal membrane disintegrates and the
enzymes are freed to act on the cellular debris to digest it. Hence, lysosomes are
often referred to as suicide bags. Hence, lysosome functions as a form of digestive
system for the cell. Each lysosome is filled with large numbers of small granules
which are protein aggregates of hydrolytic (digestive) enzymes. The main
substances lysosome digests are proteins, carbohydrates, lipids.

Peroxisomes
Peroxisomes are also small membrane-bound bodies which are similar in
appearance to lysosomes. They contain catalase which causes the breakdown of
hydrogen peroxide. The peroxisomes membrane contains some peroxisome-specific
proteins that are concerned with the transport of substances into and out of the
matrix of the peroxisome. The matrix contains more than 40 enzymes and these
enzymes operate in concert with other enzymes outside the peroxidase to catalyse
reactions, including the catabolism of very long chain fatty acids.

2.3- Cell or Plasma Membrane


The membrane that surrounds the cell is called the cell membrane. It is also referred
to as the plasma membrane. The Figure shows the structure of the cell membrane.
The cell membrane is composed primarily of membrane protein and lipid and is
about 7.5 nm (75 Angstrom units) thick. They are semipermeable allowing some
substances to pass through and excluding others. The major lipids are phospholipids
and the approximate composition of the cell membrane is proteins, 55 per cent;
phospholipids, 25 per cent; cholesterol, 13 per cent; other lipids, 4 per cent; and
carbohydrates, 3 per cent. The accepted model concept of the structure of the cell
membrane is that of a fluid mosaic model. According to this concept, the lipid
30
bilayer is in form of a fluid and membrane protein mostly lipoprotein and
glycoprotein, which are loosely attached and embedded in a bilayer matrix. Figure 6
also shows globular masses floating in the lipid bilayer. These are membrane
proteins, most of which are glycoproteins. The protein components of the cell
membrane are of two main types— integral proteins and peripheral proteins.
Integral protein pass all the way through the cell membrane, whereas, peripheral
protein are attached to the outside or inside of the cell membrane. The integral
protein provide pathway through which the water soluble substance diffuses
through the extra and intracellular fluid. The peripheral protein functions almost
entirely as enzyme. The membrane lipid makes up the matrix that give the shape and
structure to the cell membrane and embedded in this matrix are the membrane
proteins. All membrane contain phospholipid and glycolipid which are amphipathic
in nature (possess two coats).

The lipids are characterised by having hydrophobic and hydrophilic ends. The
hydrophilic end of the bi-lipid molecule are repelled by water but attracted to each
other, as shown in Figure 6. Membrane lipids are almost impermeable to water and
water soluble substances such as ions, glucose, urea, etc. while lipid soluble
substances such as oxygen, carbon dioxide, alcohol can penetrate easily.
The proteins in the cell membrane carry out many functions. They serve as:
i. Pumps which are actively involved in transporting ions across the

membrane e.g. Na+ - K+pump.


ii. Carriers transporting substances down the electrochemical gradient by a
process called facilitated diffusion.
iii. Ion channels which when activated permit the passage of ions into or out of

31
the cell.
iv. Receptors that bind neurotransmitter and hormone initiating physiological
changes inside the cell.
v. Enzyme catalysing reaction at the surface of the membrane.
vi. Antigen and antibodies, the antigenic properties of the cell depend on the
protein outside.

Figure 1.1.6: Structure of the cell membrane

ITQ 2: What are Peroxisomes?


ITA 2: Peroxisomes are also small membrane-bound bodies which are similar in appearance to
lysosomes. They contain catalase which causes the breakdown of hydrogen peroxide. The
peroxisomes membrane contains some peroxisome – specific proteins that are concerned with the
transport of substances into and out of the matrix of the peroxisome. The matrix contains more
than 40 enzymes and these enzymes operate in concert with other enzymes outside the peroxidase
to catalyse reactions, including the catabolism of very long chain fatty acids.

32
3.0 Conclusion/Summary
In this section, we have focused on the cell. I explained that the cell is the basic unit
of life with many structures organised to perform different functions that keep the
cell alive. In this study session, I am sure you have learnt that:
The cell has two major parts, the nucleus and the cytoplasm with different
substances all called protoplasm. The cell is made up of many structures and
substances that perform diverse functions. The important parts of the cell that
perform different functions include the Nucleus, the Cytoplasm, the
Endoplasmic reticulum and ribosomes, the Golgi apparatus, the Mitochondria,
Centrosome, the Lysosomes to mention a few. You have also learnt that the
cell or plasma membrane made up of protein and lipid performs about six
listed functions.

4.0 Self-Assessment Questions


1. Check this https://www.youtube.com/watch?v=kV1r2oVlHLI
2. Checking Youtube, pick the video that best helps you learn about the cell and
its functions and share the information with your colleagues in the discussion
forum online.
3. Explain the Organisation of the cell
4. Explain 3 different substances that make up the cell
5. Give detailed explanation of the Cell Structure
6. Draw a typical cell showing the organelles in the cytoplasm and the nucleus.

33
7.0 References/Further Readings

Fox SI. (2012). Human Physiology. 12th edition, Mc Graw Hill, New York.

Ganong, W.F. (2010). Review of Medical Physiology. 23rd edition, Mc


Graw Hill, New York.
Guyton, A.C, Hall J.E. (2001). Textbook of Medical Physiology. Harcourt

International Edition , 10th edition, W.B. Saunders, Philadelphia.


Oyebola DO. (2002). Essential Physiology, Vol 1, Nihort Press

34
STUDY SESSION 2
Transport Across Cell Membrane
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Simple diffusion
2.2 Facilitated diffusion
2.3 Active transport
2.4 Secondary active transport
2.5 Osmosis
2.6 Endocytosis
2.7 Exocytosis
2.8 Solvent drag-
3.0Conclusion/Summary
5.0 Self-Assessment Questions
6.0 Additional Activities (Videos, Animations & out of Class activities)
7.0 References/Further Reading

Introduction:
Let me open this session by saying that no cell is an island and that life is only
possible because several cells that make up the various organs of the body
communicate. There is movement of substances across the cells and this is
facilitated through various media. In this unit, you are going to learn about how
solutes and solvents are transported across the cells.

35
1.0 Study Session Learning Outcomes
After studying this study session, I expect you to be able to explain the following
concepts in detail:
1. Simple diffusion
2. Facilitated diffusion
3. Active transport
4. Secondary active transport
5. Osmosis
6. Endocytosis
7. Exocytosis
8. Solvent drag

2.0 Main Content


2.1. Simple diffusion
As I am sure you already know, diffusion is the movement of the molecules of a
substance from a region of higher concentration to that of lower concentration. This
movement continues until the molecules are evenly distributed in the two regions,
as demonstrated in Figure 1-9. It involves the movement of substances down their
concentration gradient. It is a passive process; that is, it does not require energy. It
is not carrier mediated. It does not display inhibition, either competitive or not. It is
not saturable.

36
Figure1.2.1: Movement of molecules by simple diffusion

ITQ 1: What is simple diffusion?


ITA 1: This is the movement of the molecules of a substance from a region of higher
concentration to that of lower concentration.

2.2 Facilitated diffusion


This is a carrier mediated transport and it involves some transport proteins that
transport substances of larger molecules across the cell membrane. (Figure 1-10). It
transports substances down their concentration gradients, thus it also is a passive
process. It is saturable and exhibits the characteristics of inhibition either
competitive or non-competitive. A typical example of facilitated diffusion is the
glucose transport by glucose transporters which move glucose down the
concentration gradient from extracellular fluid into cytoplasm of the
cell.

37
Figure1.2.2: Movement of molecules by facilitated diffusion using carrier protein

2.3 Active transport


As the name implies it is an active process that requires energy. It transports
substances against the concentration gradient. It is also carrier mediated and
saturable. It exhibits the characteristics of inhibition either competitive or non-
competitive. Energy that is utilised for this transport is obtained from adenosine
triphosphate (ATP) hydrolysis. A typical example of active transport is the
transport of sodium ion out of the cell against its concentration gradient and
the active transport of potassium ion into the cell against its own concentration

gradient by the Na+ - K+ pump (Na+- K+ ATPase). For every, Na+ pumped out by

this pump, 2K+ is pump in.

Figure 1–11 shows the basic physical components of the Na+-K+ pump. The pump
consists of a carrier protein which is a complex of two separate globular proteins, a

38
larger one having a molecular weight of 100, 000 and the small one has molecular
weight of 45, 000. Though, the function of the smaller one is not known. The larger
protein has specific features that are very important for the pump.

Figure 1. 2.3: Mechanism of the sodium-potassium pump. ADP, adenosine diphosphate;


ATP, adenosine triphosphate; Pi, phosphate ion.

2.4 Secondary Active Transport


Some transport protein are uniport because they transport only one substance,
others are called symport because the transport requires the binding of more than
one substance to the transport protein and then the substances are transported across
the membrane. An example of symport is a carrier protein in the intestinal mucosa
that is responsible for the co-transport of sodium ion and glucose from the intestinal

39
mucosa into the mucosa cells, other transporter are called antiport because they

exchange one substance for the other. The Na+ - K+ ATPase is a typical example

of an antiport. It moves three Na+ out of the cell in exchange for each two K+ that
is moved into the cell.

Figure 1.2.4: Classes of transport protein

Secondary active transport is a carrier mediated transport that involves the binding
of two types of substance to the binding site of the carrier. The carrier then
transports both substances in or out of the cell as the case maybe. One of these
substances is transported down its concentration gradient, while the other is
transported against its concentration gradient. The latter substance is being dragged
along by the former substance as it moves down its concentration gradient. The
driving force for this type of transport is supply by the concentration gradient of
one of the substances but not by ATP. This type of transport is also called Na co-
transport.

40
Glucose and many amino acids are transported into most cells against large
concentration gradients; the mechanism of this is entirely by co-transport, as shown
in Figure 1–13. Sodium co-transport of glucose and amino acid occur especially in
the epithelial cell of the intestinal tract and renal tubule to aid in the absorption of
these substances into the blood. A typical example of sodium co-transport is
demonstrated in the transport of glucose into the epithelial cells lining the small
intestine by a synport. Present in the luminal brush border membrane of the small
intestine is a synport that transport glucose into the cell following the binding of

Na+ to that carrier. The Na+ is transported down its electrochemical gradient while
the glucose is transported against its concentration gradient. The electrochemical
gradient of sodium provides the driving force for the transport of glucose
molecules. Thus, the sodium drags the glucose to transport across the brush border

membrane. The electrochemical gradient of sodium is maintained by the Na+ - K+


pump.

Figure 1.2.5: Postulated mechanism for sodium co-transport of glucose.

41
2.4 Osmosis
This is a process of diffusion of solvent molecule from a region where there is
higher concentration (low solute concentration) to a region where there is lower
concentration (higher solute concentration) across a semi-permeable or selectively
permeable membrane. To give an example of osmosis, let us assume the conditions
shown in Figures 1-14 & 1-15, with pure water on one side of the cell membrane
and a solution of sugar and sodium chloride on the other side. When a substance is
dissolved in water the concentration of water molecule in the solution is less than
that of pure water of equal volume. If the solution is placed on one side of the
membrane that is permeable to water and not to solute and an equal volume of water
is placed on the other side, water molecule diffuse down their concentration gradient
into the solution. This process of diffusion of solvent molecule to a region in which
there is a higher concentration of solute to which the membrane is impermeable is
called osmosis.

Figure 1.2.6: Demonstration of osmotic pressure caused by osmosis at a semi permeable


membrane

42
Figure 1.2.7: Osmosis at a cell membrane when a sodium chloride solution is placed
on one side of the membrane and water is placed on the other side.

The tendency for movement of solvent molecule to a region of a greater solute


concentration can be prevented by applying a pressure to the mole concentration
solution. The pressure necessary to prevent solvent migration is the osmotic
pressure of the solution. The osmotic pressure depends upon the number rather
than the type of particle insolution.

The term tonicity is used to describe the osmolality of solution relating to plasma,
solutions that have the same osmolality as plasma are said to be isotonic, those
with higher osmolality are called hypertonic, while those with lesser osmolality are
said to be hypotonic. (Figure 1-16).

43
Figure 1.2.8: Effects of isotonic, hypertonic, and hypotonic solutions on cell volume.

2.5 Exocytosis
This is the process of extrusion of substances out of the cell. Proteins that are
secreted by the cell moves from the endoplasmic reticulum to the golgi apparatus,
and from the trans-golgi they are extrude into secretory granules or vesicles. These
granules move into cell membrane. Their membrane then fuses with the cell
membrane and the area of fusion then breaks down. This leaves the content of the
granules or vesicle outside the cell while the cell membrane remains intact. This
extrusion process required calcium ion and energy.

2.6 Endocytosis
This is the process of ingestion of substances by the cell. It is the reverse of

44
exocytosis. There are two types of endocytosis, these include phagocytosis and
pinocytosis. Phagocytosis (cell eating) is the process by which bacteria, dead tissue
or other particles visible under the microscope are engulfed by cells such as the
polymorphonuclear leukocytes. The material makes contact with the cell membrane
which then invaginates, leaving the engulfed material in the membrane enclosed
vacuole while cell membrane remains intact.
Phagocytosis occurs in the following steps:
i. The cell membrane receptors attach to the surface ligands of the
particle.
ii. The edges of the membrane around the points of attachment evaginate
outward within a fraction of a second to surround the entire particle;
then, progressively more and more membrane receptors attach to the
particle ligands.
iii. Actin and other contractile fibrils in the cytoplasm surround the
phagocytic vesicle and contract around its outer edge, pushing the
vesicle to the interior.
iv. The contractile proteins then pinch the stem of the vesicle so
completely that the vesicle separates from the cell membrane, leaving
the vesicle in the cell interior in the same way that pinocytotic vesicles
are formed.
Pinocytosis means ingestion of minute particles that form vesicles of extracellular
fluid and particulate constituents inside the cell cytoplasm. Pinocytosis is
essentially the same process like phagocytosis, the only difference begin that the
substances ingested are in solution and hence not visible under the microscope.
Pinocytosis is the only means by which most large macromolecules, such as most
protein molecules, can enter cells.
45
Figure 1–17 demonstrates the successive steps of pinocytosis, showing three
molecules of protein attaching to the membrane. These molecules usually attach to
specialised protein receptors on the surface of the membrane that are specific for
the type of protein that is to be absorbed. The receptors generally are concentrated
in small pits on the outer surface of the cell membrane, called coated pits. On the
inside of the cell membrane beneath these pits is a latticework of fibrillar protein
called clathrin, as well as other proteins, perhaps including contractile filaments of
actin and myosin. Once the protein molecules have bound with the receptors, the
surface properties of the local membrane change in such a way that the entire pit
invaginates inward, and the fibrillar proteins surrounding the invaginating pit cause
its borders to close over the attached proteins as well as over a small amount of
extracellular fluid. Immediately thereafter, the invaginated portion of the
membrane breaks away from the surface of the cell, forming a pinocytotic vesicle
inside the cytoplasm of the cell.

Figure 1.2.9: Mechanism of pinocytosis

46
Solvent drag
When solvent is moving in one direction (bulk flow), the solvent tends to drag along
some molecules or solutes in that direction, this force is called solvent drag. In most
situations in the body, its effects are very small.

ITQ 2: The process of diffusion of solvent molecule to a region in which there is a higher
concentration of solute to which the membrane is impermeable is called…..
ITA 2: Osmosis

3.0 Conclusion/Summary
Transportation of materials across cells is made possible through diffusion, osmosis
and active transport using different media. In this unit you have learnt that the
various mechanisms utilized by the body include simple diffusion, facilitated
diffusion, active transport, secondary active transport, osmosis, endocytosis,
exocytosis and solvent drag. The various mechanisms allow for exchange of fluid,
minerals and molecules across cells.

4.0 Self-Assessment Questions


1. check these: https://www.youtube.com/watch?v=w3_8FSrqc-I,
https://www.youtube.com/watch?v=U9ZfowGuLfk
https://www.youtube.com/watch?v=zuNMVzTeCtw,
https://www.youtube.com/watch?v=mzo_B5F7pk4,
http://www.differencebetween.net/science/difference-between-exocytosis-and-
endocytosis/
http://www.differencebetween.net/science/difference-between-pinocytosis-and-
phagocytosis/https://www.youtube.com/watch?v=SSS3EtKAzYc

47
Do a summary of what you have learnt from each of the listed sites and submit along
with your assignments at the end of this course.
2. Distinguish between simple and facilitated diffusion
3. Explain the Sodium-potassium pump
4. Explain facilitated diffusion
5. Describe active transportation with particular reference to Mechanism of the
sodium-potassium pump

7.0 References/Further Readings

Fox SI. (2012). Human Physiology. 12th edition, Mc Graw Hill, New York.

Ganong, W.F. (2010). Review of Medical Physiology. 23rd edition, Mc


Graw Hill, New York.
Guyton, A.C, Hall J.E. (2001). Textbook of Medical Physiology. Harcourt

International Edition , 10th edition, W.B. Saunders, Philadelphia.


Oyebola DO. (2002). Essential Physiology, Vol 1, Nihort Press.

48
STUDY SESSION 3
Biologically Important Molecules and Their Functions
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Carbohydrates
2.2 Proteins
2.3 Lipids
2.4 Nucleic Acids
3.0 Conclusion/Summary
4.0 Self-Assessment Questions
5.0 Additional Activities (Videos, Animations & out of Class activities)
6.0 References/Further Readings

Introduction:
Our body needs digested nutrients to provide energy needed to perform functions
and amino acids for body building and repairs. The nutrients in various forms are
transported as small molecules of complex organic chemicals either as broken down
carbohydrates, proteins, lipids and other forms. In this unit you will be introduced
to the small molecules of these nutrients in the forms that they are absorbed.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Describe carbohydrates
2. Explain the three main classes of carbohydrates
49
3. Discuss the major types of Proteins
4. Describe lipids
5. Describe Nucleic acids

2.0 Main Content


2.1 Carbohydrates
Carbohydrates are organic compounds made of carbon, hydrogen, and oxygen
atoms. Carbohydrate are made of monosaccharide (simple sugars molecules) linked
together. Their function is to provide a key source of energy for cells. An example
is starch, made of many linked glucose molecules. Carbohydrates are divided into
three main classes: (i) monosaccharides (ii) disaccharides (iii) polysaccharides.

Monosaccharides
Monosaccharides are simple sugar unit with a general formula (CH2O)n. The ‘n’
ranges between 3 and 9. Monosaccharides are all sweet, small crystalline
molecules. They are readily soluble in water and are all reducing sugars. They are
classified on the basis of the number of carbon atoms: trioses (3 carbons), tetroses
(4 carbon), pentoses (5 carbons). The most common are pentoses and hexoses. Most
monosaccharides are metabolic energy sources and serve as building blocks for the
synthesis of other macromolecules.

Disaccharides
They are formed when two monosaccharides combine by condensation. The bonds
formed between the two monosaccharides units as a result of condensation is called
a glycosidic bond. There are several examples of disaccharide, but the most
common are maltose, lactose and sucrose.

50
Polysaccharides
These are made by joining several monosaccharide units. They function mainly as
food and energy stores e.g. starch and glycogen or as structural material e.g.
cellulose. They are not sweet, non-crystalline, either slightly or insoluble in water.

ITQ 1: What are carbohydrates?


ITA 1: Carbohydrates are organic compounds made of carbon, hydrogen, and oxygen
atoms.

2.2 Proteins
Proteins are macromolecules with molecular weight of several thousands. They are
compounds containing carbon, hydrogen, oxygen, nitrogen, sulphur. There are two
distinct types of protein:
i. Fibrous proteins
ii. Globular proteins.

Fibrous proteins are insoluble in water and are physically tough. This property
enables them to play a structural role in a cell. Major examples of fibrous proteins
are:
(a) Collagen: This is found in bones, skin, tendon and cartilage. This is

the most abundant protein in invertebrates and it usually contains three


very long polypeptide chains, each with about 1,000 amino acids.
(b) Keratin: This is found in the outermost layer of the skin and hair,

scales, hooves, nails and the feathers of animals. The main function is
to protect the body against the environment.
(c) Fibrinogen: This is blood plasma protein, responsible for blood

clotting. With the action of thrombin, fibrinogen is converted into


51
molecules of insoluble protein called fibrin, which forms a network
on the surface of wounds to trap blood cells and form clots.

Globular proteins
These are proteins that are soluble in water. They have tertiary and sometimes
quaternary structures. They are folded into spherical or globular shapes. They
include immunoglobulins or antibodies in the blood, enzymes and some hormones,
which are important in maintaining the structure of the cytoplasm.

2.3 Lipids
Lipids are non-polar molecules that are not soluble in water. They include fats,
phospholipids, steroids, and waxes. Lipids functions are to provide energy and
serve an important part in the structure and functioning of cell membranes. Some
examples of lipids include butter (saturated fat), cholesterol (steroid) and ear wax
(wax).

2.4 Nucleicacids
Nucleic acids are long chains of smaller molecules called nucleotides. Nucleic acids
mainly serve the purpose of providing the organism with its genetic blueprint and
coding. Deoxyribonucleic acid (DNA) and ribonucleic acid (RNA) are two types of
nucleic acids.

ITQ 2: Define Proteins


ITA 2: Proteins are macromolecules with molecular weight of several thousands. They are
compounds containing carbon, hydrogen, oxygen, nitrogen and sulphur.

52
3.0 Conclusion/Summary
In this unit, we have learnt that carbohydrates are made of carbon, hydrogen and
oxygen atoms metabolic energy sources and serve as building blocks for the
synthesis of other macromolecules. Protein on the other hand are made up of carbon,
hydrogen, oxygen, nitrogen, sulphur while lipids are not water soluble materials that
include fats, phospholipids, steroids, and waxes. Nucleic acid provides basis for the
genetic blueprint and coding of the DNA and RNA.

In this unit, we have learnt that carbohydrates made of carbon, hydrogen and
oxygen atoms metabolic energy sources and serve as building blocks for the
synthesis of other macromolecules. Protein on the other hand are made up of
carbon, hydrogen, oxygen, nitrogen, sulphur while lipids are not water soluble
materials that include fats, phospholipids, steroids, and waxes. Nucleic acid
provides basis for the genetic blueprint and coding of the DNA and RNA.
In this unit, you have learnt that:
i. Carbohydrates are made up of three classes of monosacharides,
disaccharides and polysaccharides depending on the number carbon
chains
ii. Proteins are mainly two types, fibrous and globular proteins
iii. Lipids are non-polar molecules, insoluble in water and include fats,
phospholipids, steroids and waxes and examples include saturated fats
and cholesterol.
iv. Nucleic Acids

53
4.0 Self-Assessment Questions
Please answer the following questions:
1. Differentiate between the three main classes of carbohydrates
2. How is fibrous protein different from globular protein
3. List two forms of lipids
4. Discuss the two distinct type of protein

5.0 References/Further Readings

Fox SI. (2012). Human Physiology. 12th edition, Mc Graw Hill, New York. Ganong,

W.F. (2010). Review of Medical Physiology. 23rd edition, Mc Graw Hill,


New York.
Guyton, A.C, Hall J.E. (2001). Textbook of Medical Physiology. Harcourt

International Edition , 10th edition, W.B. Saunders, Philadelphia.


Oyebola DO. (2002). Essential Physiology, Vol 1, Nihort Press.

54
STUDY SESSION 4
Homeostasis
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Homeostasis
2.2 The body systems
2.3 Feedback control systems
3.0 Conclusion/Summary
4.0 Self-Assessment Questions
5.0 Additional Activities (Videos, Animations & Out of Class activities)
6.0 References/Further Readings

Introduction:
The human body has a remarkable capacity for self-restoration. The Greek
physician Hippocrates (father of naturalism and rationalism) commented that the
human body usually returns to a state of equilibrium by itself and people recover
from most illnesses even without the help of a physician. This tendency results from
the body’s ability to detect change and activate mechanisms that oppose it. In this
unit, you will learn about the concept of homeostasis, how the body systems
achieve physiological body maintenance through feedback mechanisms.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Define Homeostasis
55
2. Explain the different types of body systems that contribute to
homeostasis
3. Explain the different types of feedback control systems that the body
uses to maintain stability.

2.0 Main Content


2.1 Homeostasis
Homeostasis is the maintenance of a relatively constant internal environment in an
ever changing external environment. It comes from the word homeo, which means
the sameness, and stasis, that is, standing still. We can also define it as the
maintenance of the constancy of the composition of the internal environment. The
mechanisms which work towards its achievement are called homeostatic
mechanisms. Essentially all the organs and tissues of the body perform functions
that help to maintain these constant conditions. For instance, the lungs provide
oxygen to the extracellular fluid to continually replenish the oxygen that is being
used by the cells, the kidneys maintain constant ion concentrations, and the
gastrointestinal system provides nutrients.

Each body system contributes to the homeostasis of other systems and of the entire
being. No system of our body works in isolation, and our well-being depends upon
the well-being of all the interacting body systems. A disruption within one system
generally has consequences for several additional body systems. Let us consider
some brief explanations of how various body systems contribute to the maintenance
of homeostasis.

56
ITQ 1: Define homeostasis
ITA 1: Homeostasis is the maintenance of a relatively constant internal environment in an
ever changing external environment.

2.2. The Body Systems and Their Contributions to Homeostasis.


Nervous system
This system is made up of the brain, spinal cord, nerves and receptors. The nervous
system, along with the endocrine system, serves as the primary control centre of our
body. It operates at a subconscious level and controls many functions of the internal
organs, including the level of pumping activity by the heart, movements of the
gastrointestinal tract, and secretion by many of the body’s glands. For example, the
hypothalamus of the brain is where the body's "thermostat" is found. The
hypothalamus also stimulates the pituitary gland to release various hormones that
control metabolism and development of the body. The sympathetic and
parasympathetic divisions of the nervous system alternatively stimulate or inhibit
various bodily responses (such as heart rate, breathing rate, etc.) to help maintain
them at optimum levels.

It also controls contraction of muscles like the erector pili muscles (involved in
thermoregulation) and skeletal muscles. The nervous system also regulates various
systems such as the respiratory (controls rate and depth of breathing),
cardiovascular system (controls heart rate and blood pressure), endocrine organs
(causes secretion of antidiuretic hormone ADH and oxytocin), the digestive system
(regulates the digestive tract movement and secretion), and the urinary system
(helps adjust renal blood flow and also controls voiding the bladder). The nervous
system is also involved in sexual behaviours and functions.

57
2.3 Endocrine system
The endocrine system consists of glands hypothalamus, pituitary, thyroid, adrenal
testes and ovaries which secrete hormones into the bloodstream. Each hormone has
an effect on one or more target tissues. In this way the endocrine system regulates
the metabolism and development of most body cells and body systems. Bone
growth is regulated by several hormones, and the endocrine system helps with the
mobilisation of calcium and phosphate into and out of the bones. In the muscular
system hormones adjust muscle metabolism, energy production, and growth. In the
nervous system, hormones affect neural metabolism, regulate fluid/electrolyte
balance and help with reproductive hormones that influence central nervous system
(CNS) development and behaviours. In the cardiovascular system hormones are
needed in the regulation of RBC's production, and blood pressure. Hormones also
have anti-inflammatory effects as well as stimulate the lymphatic system. In
summary, the endocrine system has a regulatory effect on basically every other
body systems.

Skeletal system
It consists of all bones in the body, cartilages and ligaments. The skeletal system
serves as an important mineral reserve. For example, if blood levels of calcium or
magnesium are low and the minerals are not available in the diet, they will be taken
from the bones. On the other hand the skeletal system provides calcium needed for
all muscle contractions. Lymphocytes and other cells relating to the immune
response are produced and stored in the bone marrow. The skeletal system aids in
protection of the nervous system, endocrine organs, chest and pelvic regions in
which vital organs are housed.

58
Integumentary system
This system is composed of the skin that is the epidermis, dermis and adipose
tissue, nails, hair, receptors, oil glands and sweat glands. The integumentary system
is involved in protecting the body from invading microbes, regulating body
temperature through sweating and vasodilation, or shivering and piloerection, and
regulating ions balance in the blood. Stimulation of mast cells also produces
changes in diameter of blood vessels and capillary permeability which can affect
the blood flow in the body and how it is regulated. It also helps synthesise vitamin D
which interacts with calcium and phosphorus absorption, a factor that is very
important for bone growth and maintenance. Hair on the skin guards entrance into
the nasal cavity or other orifices preventing invaders from getting further into the
body. The skin also helps maintain balance by excretion of water and other solutes.
The keratinised epidermis limits fluid loss through the skin, thus providing
mechanical protection against environmental hazards.

Lymphatic system
The lymphatic system is composed mainly of the lymphatic vessels, lymph nodes,
thymus, spleen and the bone marrow. It has three principal roles. First is the
maintenance of blood and other body fluid volumes. Excess fluid that leaves the
capillaries when under pressure would build up and cause edema, but for the role of
the lymphatic system. Secondly, the lymphatic system absorbs fatty acids and
triglycerides from fat digestion so that these components of digestion do not enter
directly into the blood stream. Thirdly, the lymphatic system is involved in
defending the body against invading microbes, and also in the immune response.
This system assists in body maintenance such as bone and muscle repair after
injuries. It also assists in maintaining the acid pH of urine required to fight

59
infections in the urinary system. The tonsils are the body helpers that defend against
infections and toxins absorbed from the digestive tract. The tonsils also protect
against infections entering into the lungs.

Respiratory system
The components of the respiratory system are the nasal cavity, pharynx, larynx,
glottis, epiglottis, bronchi, bronchioles, alveoli and the lungs. The respiratory
system works in conjunction with the cardiovascular system to provide oxygen to
cells within every body system for cellular metabolism. The respiratory system also
removes carbon dioxide. Since CO2 is mainly transported in the plasma as
bicarbonate ions, which act as a chemical buffer, the respiratory system also helps
maintain proper blood pH levels, a fact that is very important for homeostasis. As a
result of hyperventilation, the level of CO2 is reduced. This causes the pH of body
fluids to increase. If pH rises above 7.45, the results are respiratory alkalosis. On
the other hand, too much CO2 causes pH to fall below 7.35 which results in
respiratory acidosis. The respiratory system also helps the lymphatic system by
trapping pathogens and protecting deeper tissues from invading microorganisms.

Urinary system
Its main components are the kidneys, ureter, bladder and urethra. Toxic nitrogenous
wastes cumulate as urea, uric acid and creatinine. The urinary system rids the body
of these wastes. It is also involved in the maintenance of blood volume, blood
pressure and electrolyte concentrations within the blood. The kidneys produce a
hormone (erythropoietin) that stimulates red blood cell production. They also play
an important role in maintaining the water content of the body and the level of salts
in the extracellular fluid.

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Cardiovascular system
It consists of the heart, blood vessels and the blood. The cardiovascular system
ensures the normal functioning of other body systems by transporting hormones,
oxygen and nutrients to them and taking away waste products from them thereby
providing all living body cells with a fresh supply of oxygen and nutrients and also
removing carbon dioxide and other toxic wastes from their surroundings.
Homeostasis is disturbed if the cardiovascular or lymphatic systems are not
functioning properly. The cardiovascular system also contains sensors to monitor
blood pressure. They are called baroreceptors. They detect the amount of stretch of
the blood vessels and relay information via the nerves to the CNS which brings
about the appropriate responses that regulate the blood pressure.

Muscular system
This system is made of skeletal muscles such as biceps, quadriceps, and
gastrocnemius muscles and smooth or involuntary muscles such as cardiac muscle,
intestinal muscles and muscles of the blood vessels. The muscular system is largely
responsible for movement, posture, balance, gait, secretion by glands and
maintenance of body temperature through heat production. It also contributes to
blood glucose balance by storing glucose in form of glycogen. Muscles also aid in
moving blood through veins, protect deep blood vessels and help the lymphatic
system move lymph.

Digestive system
Its components include oral cavity, esophagus, stomach, intestines, liver and
pancreas. The nutrients needed by the body are derived from the diet. Food is taken
in by the mouth and broken down into its component parts by enzymes in the
gastrointestinal tract (or gut). The digestive products are then absorbed into the
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blood across the wall of the intestine and pass to the liver via the portal vein. The
digestive system absorbs organic substances, vitamins, ions, and water that are
needed all over the body. The liver makes nutrients available to the tissues both for
their growth and repair and for the production of energy.

Reproductive system
The main components of this system are the ovaries, testes, prostate, uterine tubes,
uterus, vagina and penis. The reproductive system is responsible for the production
of sperm cells and oval for the production of new offspring. The sex hormones do
have various effects on other body systems, and an imbalance can lead to various
disorders.

Feedback systems: negative feedback control


Negative feedback is the mechanism by which our body maintains conditions
within particular limits. It is a control system that acts to maintain the level of some
variable within a given range following a disturbance. Once equilibrium conditions
are restored, the stimulus that activated the feedback loop is removed, so that the
system ceases to function until an appropriate stimulus initiates the feedback
process again; that is, negative feedback systems in the body normally are
reversible and they come into play on demand. The component of a simple negative
feedback loop include (i) a regulated variable, (ii) sensor (or detector), (iii)
controller (comparator), and (iv) Effector. Each component controls the next
component to it (Figure 1.1).

Various disturbances may arise within or outside the internal environment and
cause undesirable changes in the regulated variable. The regulated variable is

62
sensed by sensor, information about its level is fed back to a controller
(comparator), which compares it to a desired value (set point). If there is a
difference, signal is generated, which drives the effector to oppose the changes and
bring the regulated variable closer to the desired position.

Figure 1.4.1: Component of a simple negative feedback loop

A familiar example of a negative feedback control is the thermostatic control of


room temperature. Room temperature (regulated variable) is subject to disturbance;
on a cold day, room temperature falls. The room temperature is detected by a
thermometer (sensor) in the thermostat (controller). The thermostat is set for a
certain temperature (set point). The controller compares the actual temperature
(feedback signal) to the set point temperature and signal is generated if the former

63
falls below the latter. The signal activates the furnace (effector). The resulting
change in temperature is monitored by the controller, and when temperature rises
sufficiently the furnace is turned off. Such a negative feedback system allows some
fluctuation in room temperature. Effective communication between the sensor and
effector is important in keeping these oscillations to a minimum.

Similar negative feedback systems maintain homeostasis in our body. One example
is in arterial blood pressure regulation illustrated in Figure 1.2. These system
sensors (arterial baroreceptors) are located in the carotid sinuses and aortic arch.
Changes in stretch of the walls of the carotid sinus and aorta, which follow from
changes in blood pressure, stimulate these sensors. Afferent nerve fibres transmit
impulses to control centres in the medulla oblongata. Efferent nerve fibres send
impulses from the medullar centre to the systems effectors, the heart and blood
vessels. The output of blood by the heart and resistance to blood flow are altered in
an appropriate direction to maintain blood pressure, as measured at the sensors
within a given range values.

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Figure 1.4.2: Regulation of arterial blood pressure

The control of testosterone secretion, control of calcium ions level in the blood,
control of blood glucose by insulin and glucagon, control of cortisol secretion by
the adrenal cortex are other examples of the operation of such mechanisms.

Positive feedback control


Positive feedback is a self-amplifying cycle in which a physiological change leads
to even greater changes in the same direction, rather than producing the corrective

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effects of negative feedback. Positive feedback promotes rapid change and it is
often a normal way of producing rapid progressive change in one direction. For
example, when a woman is giving birth, the head of the baby pushes against her
cervix and stimulates nerve endings there. Nerve signals are sent to the brain,
which, in turn, stimulates the pituitary gland to secrete the hormone oxytocin.
Oxytocin travels in the blood and stimulates the uterus to contract. This pushes the
baby downward, stimulating the cervix the more and causing the positive feedback
loop to be repeated. Labour contractions therefore become more and more intense
until the baby is expelled.

Figure 1.4.3: Control of childbirth by positive feedback mechanism


It should be noted however that the overall process of childbirth is a negative
feedback loop— it is a response to pregnancy that terminates the pregnancy. But
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within this negative feedback loop, there is a smaller positive feedback loop that has
just been described. Beneficial positive feedback loops are often part of larger
negative feedback loops. Other examples of beneficial positive feedback includes:
generation of nerve signals, blood clotting and the stomach digestion of protein.
Frequently, however, positive feedback is a harmful and even life-threatening
process. This is because its self-amplifying nature can quickly change the internal
state of the body to something far from its homeostatic set point. Consider a high

fever, for example. A fever, but if the body temperature rises much above 42oC, it
may create a dangerous positive feedback loop. This high temperature raises the
metabolic rate, which makes the body to produce heat faster than it gets rid of it.
Thus temperature rises still further, increasing the metabolic rate and heat
production still more.

This “vicious circle” becomes fatal at approximately 45oc. Such a temperature is so


high that they destroy the proteins that cells need to function. Convulsion and coma
are some outward signs of this damage. Thus positive feedback loops often create
dangerously out of control situations that require emergency medical treatment.

Feed forward control


Feed forward control is another strategy used to control systems in our body,
particularly when a change with time is desired. It is anticipatory in nature. A feed
forward controller generates commands without directly sensing the regulated
variable. These commands specify the target or goals. Feed forward control often
senses a disturbance and can therefore take corrective action that anticipates
change. It often operates through the feedback controllers. The moment-to-moment
operation of the feed forward controller is “open loop” (unlike closed loop in
negative feedback) because the regulated variable itself is not sensed by sensor.
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Examples include increased heart rate and breathing rate even before we have
begun to exercise, flight reactions and others.

ITQ 2: What is Positive Feedback Mechanism?


ITA 2: Positive feedback is a self-amplifying cycle in which a physiological change leads to even
greater changes in the same direction, rather than producing the corrective effects of negative
feedback. Positive feedback promotes rapid change and it is often a normal way of producing
rapid progressive change in one direction.

3.0 Conclusion/Summary
In this session, we have looked at how the principle of homeostasis allows our body
to maintain a state of balance. All the systems are involved in the complicated
process of maintaining constancy. Homeostatic control is achieved within a complex
process involving the receptor, the control centre and the effector. Homeostatic
imbalance results to diseases.

I am sure you have also learnt that:

I. Homeostasis as the ability of the body to maintain relatively stable internal


conditions even when the outside environment changes on a continuous
basis.
II. All the body systems are involved in the process of attainment of
homeostasis.
III. The body uses negative and positive feedbacks and the feed forward control
of regulating homeostatic processes in the body.

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4.0 Self-Assessment Questions
1. Watch this video clips:
https://www.youtube.com/watch?v=XZxuQo3ylIIhttps://www.youtu
be.com/watch?v=IoU3lKrOYMY
Explore the use of thirst and sweat in achieving body’s homeostasis, Explore 5
other actions of the body that contribute to the maintenance of the body and how
you can use them as guides in providing nursing care. Submit your findings to the
tutor 2 weeks after the completion of this unit.
2. Define homeostasis, and identify the components of negative feedback loops.

3. How do negative and positive feedbacks help to maintain the body homeostasis?
Illustrate these with drawing and labelling of examples of negative and positive
feedback?

7.0 References/Further Readings

Fox SI. (2012). Human Physiology. 12th edition, Mc Graw Hill, New York. Ganong,

W.F. (2010). Review of Medical Physiology. 23rd edition, McGraw Hill, New York.
Guyton, A.C, Hall J.E. (2001). Textbook of Medical Physiology. Harcourt

International Edition , 10th edition, W.B. Saunders, Philadelphia.


Oyebola DO. (2002). Essential Physiology, Vol 1, Nihort Press.
John Campbell Homeostasis 1 https://www.youtube.com/watch?v=5HS66q_OA8g
accessed on June 30 2015.
John Campbell Homeostasis 2 https://www.youtube.com/watch?v=IoU3lKrOYMY
accessed on June 30, 2015

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MODULE 2
Nerve and Muscle Physiology, Blood Physiology (I)
-Haemodynamics and Immunology
Contents
Study Session 1: Nerve and Muscle Physiology
Study Session 2: Body Fluids
Study Session 3: Haemostasis.
Study session 4: Immune System

STUDY SESSION 1
Nerve and Muscle Physiology
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Nerves
2.2 Muscle Contraction
3.0Conclusion/Summary
4.0 Self-Assessment Questions and Answers
5.0 Additional Activities (Videos, Animations & out of Class activities)
6.0 References/Further Readings

Introduction:
We respond to our living environment through our senses. We are only able to do
this because we have a master controlling and communicating systems, the
nervous system. In this unit, you are going to learn more about the typical nerve
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cell and how the nerve cells perform their functions. You are also going to learn
about how the nerve cells enable our body to engage in coordinated movement as
it facilitates muscle contractions.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Discuss the structure and functions of a typical nerve cell.
2. Explain the functional unit of the muscle
3. Discuss the mechanisms involved in muscle contraction.

2.0 Main Content


2.1 Nerves and the Functions
Nerves/Neurons are the basic structural and functional units of the nervous system.
They are specialised to respond to physical and chemical stimuli, conduct
electrochemical impulses, and release chemical regulators. Through these activities,
neurons enable the perception of sensory stimuli, learning, memory, and the control
of muscles and glands. Neurons have three principal regions: cell body, dendrites,
and axon. They vary considerably in size and shape.

Figure 2.1. 1: Parts of a neuron (myelinated)


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Figure 2.1.2: Parts of a neuron (unmyelinated)

The cell body (Soma) is the enlarged portion of the neuron that contains the
nucleus. Dendrites are thin, branched processes that extend from the cytoplasm of
the cell body. Dendrites provide a receptive area that transmits graded
electrochemical impulses to the cell body. The axon is a longer process that
conducts impulses, called action potentials, away from the cell body. The origin of
the axon near the cell body is an expanded region called the axon hillock; it is here
that action potentials originate. Side branches called axon collaterals may extend
from the axon. The axon at its end is divided into terminal branches. Each terminal
branch ends in synaptic knobs or terminal buttons. The axon of a neuron can either
be myelinated or unmyelinated. The myelinated neuron is wrapped by Schwann
cells, which form a myelin sheath (Figs 2-1 and 2-2). The myelin sheath envelops
the axon except at the terminal endings and at the Nodes of Ranvier.

Resting and Action Membrane: Resting Membrane Potential.


When the cell is not transmitting an impulse, the trans-membrane potential is called
the resting membrane potential (RMP). Also the RMP can be defined as the inside
negative potential across the membrane of cells. A resting membrane potential is
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due to uneven distribution of ions between the inside and the outside of the
membrane.
The following phenomena are involved in establishing the cell potential.
i. By means of active transport: sodium is actively pumped out of the

cell and potassium is pumped into it. So the K+ concentration in the


cell is twenty times the concentration in the extracellular fluid.

ii. The membrane at rest is far more permeable to K+ than Na+. K+ ions

diffuse out of the cell with far greater ease than Na+ diffuse into the
cell.
iii. The interior of the cell contains a high concentration of non-diffusible
ions. Of particular importance in this regard are proteins, organic
phosphates and organic sulphate anions. Since the resting membrane

is much more permeable to K+ than to Na+, the RMP is much closer

to the K equilibrium potential than that of Na+.

The chief determinants of the movement of substances across the cell membrane
are the membrane permeability, electrical as well as chemical gradients of the ions.
When the chemical and electrical forces acting on ions are equal and opposite, there
is no net flux and the system is in equilibrium.

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Action potential This is the voltage of the cell membrane when the cell membrane
is stimulated or activated. We can also define it as the potential generated when
excitable tissue (nerve and muscle) are stimulated resulting in the propagation of an
impulse. The components of the action potential are: latent period, depolarisation,
repolarisation and hyperpolarisation. (Figure 2-3).

Figure 2.1.3: Phases of action potential

When a stimulus is applied to an axon, there is a brief irregular deflection of the


baseline, called stimulus artefact. The stimulus artefact is followed by isopotential
interval (latent period) that ends with the start of action potential and corresponds
to the time it takes the impulse to travel along the axon from the site of stimulation
to the recording electrodes.

Depolarisation stage
At this time, the membrane suddenly becomes very permeable to sodium ions,
allowing tremendous numbers of positively charged sodium ions to diffuse to the
interior of the axon. The normal “polarised” state of –90 millivolts is immediately
neutralised by the inflowing of positively charged sodium ions, with the potential
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rising rapidly in the positive direction. This is what we call depolarisation. In large
nerve fibres, the great excess of positive sodium ions moving to the inside causes
the membrane potential to actually “overshoot” beyond the zero level and to
become somewhat positive. In some smaller fibres, as well as in many central
nervous system neurons, the potential merely approaches the zero level and does
not overshoot to the positive state.

Repolarisation stage
Within a few milliseconds after the membrane becomes highly permeable to
sodium ions, the sodium channels begin to close and the potassium channels open
more than normal. Then, rapid diffusion of potassium ions to the exterior re-
establishes the normal negative resting membrane potential. This is called
repolarisation of the membrane. The sharp rise and the rapid fall are the spike
potential of the axon, and the slower fall at the end of the process is the after-
depolarisation.

After the action potential, during the recovery period, Na+ that came in during

depolarisation and the K+ that went out during repolarisation are brought back to
their original positions. Since this means moving sodium against its concentration

gradient (i.e. from in to out) and vice-versa for K+, the process involves active
transport requiring energy.

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Figure 2.1.4: Characteristics of the voltage-gated sodium and potassium channels

Changes in excitability during action potential


The refractory period is divided into an absolute refractory period, which
corresponds to the period from the time the firing level is reached until
repolarisation is about 1/3 complete, and a relative refractory period, lasting from
this point to the start of after-depolarisation. During the absolute refractory period,
no stimulus, no matter how strong will excite the nerve, but during the relative
refractive period, stronger than normal stimuli can cause excitation. During after-
depolarisation, the threshold is again decreased and during after-hyperpolarisation it
is increased.

All-or-nothing principle
Once an action potential has been elicited at any point on the membrane of a normal
fibre, the depolarisation process travels over the entire membrane if conditions are
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right, or it does not travel at all if conditions are not right. This is called the all-or
nothing principle, and it applies to all normal excitable tissues.

Conduction of impulse along nerve fibres: Non-myelinated nerve


The nerve cell membrane is polarised at rest, positive charges lined up along the
outside of the membrane and negative charges along the inside of the membrane.
During the action potential (AP), this polarity is abolished and for a period is
actually reversed. Positive charges from the membrane ahead of and behind the AP
flows into the area of negativity represented by the AP (current sink). By drawing
off positive charges, this flow decreases the polarity of the membrane ahead of the
AP. This type of electrotonic depolarisation initiates a local response, and when the
firing level is reached, a propagated response occurs that in turn electrotonically
depolarises the membrane in front of it. This sequence of event moves regularly
along an unmyelinated axon to its end. Thus, the self-propagating nature of the
nerve impulse is due to circular current flow and successive electrotonic
depolarisation to the firing level of the membrane ahead of the action potential.

Myelinated nerve
Conduction in myelinated axons depends upon a similar pattern of circular current
flow. However, myelin is an effective insulator and current flow through it is
negligible. Instead, depolarisation in myelinated axons jumps from one node of
Ranvier to the next, the “current sink” at the active node serving to electrotonically
depolarize to the firing level the node ahead of the AP. This jumping of
depolarization from node to node is called saltatory conduction, as shown in Figure
2-5. It is a rapid process, and myelinated axon conducts up to 50 times faster than
the fastest unmyelinated fibres.

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Figure 2.1.5: Saltatory conduction along a myelinated axon. Flow of electrical
current from node to node is illustrated by the arrows.

Neuromuscular Transmission
As the axon supplying the skeletal muscle fibre approaches its termination, it loses
its myelin sheath and divides into a number of terminal buttons or end-feet. The
end-feet contain many small, clear vesicles that contain acetylcholine, which is the
transmitter at this junction. The endings fit into depressions in the motor end plate,
which is the thickened portion of the muscle membrane of the junctions. The
depression is called the synaptic gutter or synaptic trough, and the space between
the terminal and the end plate is called the synaptic cleft or space. At the bottom of
the gutter are numerous smaller folds of the muscle membrane of the end plate
called sub-neural clefts or functional folds, which greatly increase the surface area
which the synaptic transmitter can act on. The whole structure is known as the
neuromuscular or myoneural junction (Figure 2.6).

In the axon terminal are many mitochondria that supply ATP, the energy source that
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is used mainly for synthesis of the excitatory transmitter called acetylcholine. The
acetylcholine in turn excites muscle fibre membrane. Acetylcholine is synthesised
in the cytoplasm of the terminal button, but it is absorbed rapidly into many small
synaptic vesicles. In the synaptic space, a large quantity of the enzyme acetyl
cholinesterase, which is capable of destroying acetylcholine after it has been
released from the synaptic vesicles are present.

Figure 2.1.6: Shows neurotransmission at neuromuscular junction

ITQ 1: Three principal regions of the neuron are:


ITA 1: Cell body, dendrites, and axon.

2.2. Muscle Contraction


Three types of muscle cells can be identified on the basis of structure and contractile
properties:
(i) Skeletal muscle (ii) Smooth muscle (iii) Cardiac muscle

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Most skeletal muscles as the name implies are attached to bones and their
contraction is responsible for the movement of parts of the skeleton. Contraction of
the skeletal muscle is controlled by the somatic nervous system and hence is under
voluntary control. The movements produced by skeletal muscles are primarily
involved with interactions between our body and external environment.

Smooth muscles surround hollow organs and tubes-like stomach, intestinal tract,
urinary bladder, uterus, blood vessels and air passages to the lungs. It is also found
as single cells distributed throughout the organs (spleen) and the small group of
cells attached to the hairs in the skin.

The contraction of the smooth muscle may either propel the luminal content out of
or through the hollow organs or it may regulate the flow of the contents through
tubes by changing their diameters without itself initiating propulsion. Smooth
muscle contraction is controlled by factors intrinsic to the muscle itself by the
autonomic nervous system (ANS) and by hormones. Therefore, it is not normally
under direct conscious control. The third type of muscle; cardiac muscle is the
muscle of the heart and its contraction propels blood through the circulatory system.
Like smooth muscle, it is regulated by intrinsic factors and by ANS and hormones.

Skeletal muscle
Figure 2-7 shows the organisation of skeletal muscle. Each skeletal muscle fibre is a
cylinder with diameter of 10-100μm and length which may extend up to 300,
000μm (1 foot). The term skeletal muscle refers to a number of muscle fibres bond
together by connective tissue. From the light microscope, the most striking picture
is series of transverse light and dark bands forming a regular pattern along each

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fibre. Most skeletal and cardiac muscle fibres have these characteristic banding and
are known as striated muscles. Smooth muscle cells show no binding patterns.
Although the pattern appears to be continuous across the entire cytoplasm of a
single fibre, the bands are actually confined to a number of independent cylindrical
elements, known as myofibrils. Each myofibril is about 1 to 2 micron (μm) in
diameter and continues throughout the length of the muscle fibre. Myofibrils
occupy about 80% of the fibre volume and vary in number from several hundred to
several thousand per single fibre, depending on the fibre diameter.

The myofibrils consist of smaller filaments which are arranged in a repeating


pattern along the length of the fibril. One unit of this repeating pattern is known as a
sarcomere (little muscle), which is the functional unit of the contractile system in
striated muscles. Each sarcomere contains two types of filaments: Thick filament
composed of the contractile protein called myosin and thin filaments containing the
contractile protein components: (i) Actin (ii) Tropomyosin (iii)Troponin. Troponin
is made up of three subunits: (i) Troponin I (ii) Troponin T (iii)Troponin C. The
thick filaments, 12-18nm in diameter are located in the central region of the
sarcomere, where their orderly parallel arrangements gives rise to the dark bands
known as A-bands, because they are anisotropic to polarised light. Thin filaments,
5-8nm in diameter are attached at either end of a sarcomere to a structure known as
Z-line. Two successive Z-lines define the limits of the sarcomere. Z-lines are short
fibrous structures, which interconnect the thin filaments from two adjoining
sarcomeres thus, provide an anchoring point for the thin filaments, which extends
from the Z-lines towards the centre of the sarcomere where they overlap with the
thick filament.

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Between the ends of the dark A-bands of two adjacent sarcomeres is the I-band
(because it is isotropic to polarised light) forming the lighter region of the striated
pattern. One additional band called the H-zone appears as a thin lighter band in the
centre of the A-band. It corresponds to the space between the ends of the thin
filament. Thus, we can only find thick filaments in the H-zone. Finally, a thin dark
band can be seen in the centre of the H-zone, known as the M-line and is produced
by linkages between the thick filaments. The M-line by cross linking the thick-
filaments keeps all these in a single sarcomere in parallel alignments. Thus, neither
the thin nor thick filaments are free floating, each is linked either to Z-lines in the
case of the thin filaments or to M-lines in the case of the thick filaments.

Figure 2.1.7: Organisation of skeletal muscle from the gross to the molecular level.

Smooth muscle
Smooth muscles are distinguished anatomically from skeletal and cardiac muscles,

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because they lack physical cross striations. Actin and myosin are present and they
slide on each other to produce contraction. However, they are not arranged in
regular arrays as in skeletal muscle and cardiac muscles, and so the striations are
absent. Instead of Z-lines, there are dense bodies in the cytoplasm attached to the
cell membrane and these are bound by α – actinin to actin filament. Smooth
muscles also contain tropomyosin, but troponin appears to be absent. Smooth
muscles can be generally divided into two major types, which are shown in Figure
2-8: multiunit smooth muscle and single unit or visceral muscle.

Multiunit smooth muscle


This type of smooth muscle is composed of discrete smooth muscle fibres. Each
fibre operates entirely independently of the other fibres and is often innervated by a
single nerve ending as occurred for skeletal muscle. Furthermore, the outer surfaces
of these fibres, like those of skeletal muscle fibres, are covered by a thin layer of
glycoprotein that helps to insulate the separate fibres from each other. The most
important characteristics of multi-unit smooth muscle fibres is that their control is
exerted almost entirely by nerve fibres and very little by other stimuli, like local
tissue factors. This is in contrast to a major share of the control of visceral smooth
muscle by non-nervous stimuli. Some examples of multi-unit smooth muscle found
in the body are smooth muscle fibre of the ciliary muscle of the eye, the iris of the
eye, the nictitating membrane that covers the eye of some lower animals, the pilo-
erector muscles that cause erection of the hairs when stimulated by the sympathetic
nervous system and the smooth muscle of many of the larger blood vessels.

Visceral smooth muscle (Single unit)


Their fibres are similar to multi-unit fibres except that they are regularly or usually
83
arranged in sheet or bundles and the cell membrane contact each other at multiple
points to form many gap junctions. Thus, the fibres form a functional syncytium
that usually contracts a large area at once. For this reason, this type of smooth
muscle is also known as single unit or unitary smooth muscle. This type of muscle
is found in most of the organs in the body, especially in the walls of the gut, the bile
duct, ureters, uterus etc.

Figure 2.1.8: Multi-unit (A) and Unitary (B) smooth muscle.

Cardiac muscle
You will find that striations in cardiac muscle are similar to those in skeletal
muscle. There are large numbers of elongated mitochondria in close contacts to the
muscle myofibrils and the muscle fibres branch and interdigitate. But each is a
complete unit surrounded by a cell membrane. When the end of one muscle fibre
joins on another, the membranes of both fibres parallel each other through an
extensive series of folds. These areas which always occur as Z-lines are called
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intercalated discs. They provide a strong union between fibres, maintaining cell-cell
cohesion, so that the pull of one contractile unit can be transmitted along its axis to
the next. Along the site of the muscle fibres next to the disks, the cell membranes of
adjacent fibres fuse for considerable distances forming gap junctions. These
junctions provide low-resistance bridges for the spread of excitation from one fibre
to another.

Contractile proteins
These are proteins which participate in the contractile processes. They include
muscle proteins as well as those found in other cells and tissues. In the cells and
tissues, these proteins participate in localised contractile events in the cytoplasm, in
motile activity, and in cell aggregation phenomena. The two types of contractile
proteins that are found within muscles are actin and myosin. Both proteins are
responsible for muscle movement. The heads and necks of the myosin molecules
forms cross-links to actin.

Excitation-contraction coupling
These are the events occurring between the excitation of a muscle fibre and the
resulting contraction. The skeletal muscle fibre is so large that action potential (AP)
spreads in along the surface membrane and causes almost no current flow deep
within the fibre. However, to cause muscle contraction, this electrical current must
penetrate deeply into the muscle fibre to the vicinity of all the separate myofibrils.
This is achieved by transmission of APs along transverse tubules (T-tubules) that
penetrate all the way through the muscle fibre from one side to the other. T-tubules

action potential in turn causes release of Ca2+ in the immediate vicinity of all the

myofibrils. This Ca2+ then causes contraction. This overall process is called
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excitation contraction- coupling (Figure 2-9).

Figure 2.1.9: Excitation-contraction coupling in the muscle.

Molecular basis of contraction


Figure 2.10 demonstrates the basic mechanism of muscle contraction. This is the
process by which the shortening of the contractile elements in the muscle is brought
about by sliding of the thin filaments over the thick filaments. The width of the A-
bands is constant, whereas the Z-lines move closer when muscles contract and far
apart when it is stretched (Sliding filament mechanism). Sliding during muscle
contraction occurs when the myosin heads bind firmly to actin, bend at the junction
of the head with the neck and then detach. “This power stroke” depends on the
simultaneous hydrolysis of ATP.

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Calcium ions initiate contraction by binding troponin C. In resting muscle, troponin
I is tightly bound to actin and the tropomyosin covers the site, where myosin heads
bind to actin. Thus, the troponin-tropomyosin complex constitutes a relaxing

protein that inhibits the interaction between actin and myosin. When the Ca2+ is
released from the terminal cisterna by the AP, it binds to troponin C, the binding of
troponin I to actin is presumably weakened and this permits the tropomyosin to
move laterally. This movement uncovers the active site of myosin heads. Adenosine
triphosphate (ATP) is then split and contractions occur.

Figure 2.1.10: Relaxed and contracted states of a myofibril showing (top) sliding of the actin
filaments (pink) into: the spaces between the myosin filaments (red), and (bottom) pulling of
the Z membranes toward each other.

ITQ 2: Contraction of the skeletal muscle is controlled by the……


ITA 2: Somatic nervous system

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3.0 Conclusion/Summary

In this session, we have been able to see how nerve cells help the body to respond to
various stimuli through the mediating effect of sodium, potassium and calcium ions.
We also saw how the three main muscle types are distinguishable by their structure
and mode of contractions. I believe you have learnt about the following:
I. The structure and functions of a typical nerve cell.
II. The functional unit of the muscle
III. The various mechanisms involved in muscle contraction.

4.0 Self-Assessment Questions


As prescribed in the laboratory practical to be conducted by the Facilitator,
Answer the following questions:
1. Discuss the structure and functions of a typical nerve cell.
2. Explain the functional unit of the muscle
3. Discuss the mechanisms involved in muscle contraction.

7.0 References/Further Readings

Ganong, W. F. (2010). Review of Medical Physiology. 23rd edition, McGraw


Hill, New York.
Guyton, A.C, Hall JE. (2001). Textbook of Medical Physiology. Harcourt

International Edition. 10th edition, W.B. Saunders, Philadelphia.

Fox SI. (2012). Human Physiology. 12th edition, McGraw Hill, New York.

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STUDY SESSION 2
Body Fluid
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Body fluid Compartments
2.2 Intracellular Fluid (ICF)
2.3 Extracellular Fluid (ECF)
2.4 Measurement of Body Fluid compartments
2.4 Blood
4.0 Conclusion/Summary
5.0 Self-Assessment Questions
6.0 Additional Activities (Videos, Animations & out of Class activities)
7.0 References/Further Readings

Introduction:
The body is made of more fluids in different locations and, as a nurse, you must be
appropriately informed about the location, constituents and functions of the body
fluids. In this unit, you are going to study the body fluids in the various compartments.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Describe the different types of body fluids and where they are located.
2. Explain the method of measurement of Body Fluid in the different compartments
3. Discuss the constituents and functions of blood.
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4. Explain the concept of blood grouping and the implications in nursing practice.

2.0 Main Content


2.1 Body Fluid Compartment
General considerations and inter-relationships of body fluid spaces.
Water is an important component of the human body. It constitutes approximately
60% of body weight. In terms of volume, total body water in adult man is about 42
liters. Body fluids can be divided into two main compartments: intracellular fluid
(ICF) and extracellular fluid (ECF) in the figure below.

Figure 2.2.1 Organization of body fluids and electrolytes into compartments.

2.2 Intracellular fluid (ICF)


ICF consists of all fluid within the cell membranes of the body and is the largest
fluid compartment, accounting for 40% of the body weight. Its volume is about 28
litres. Much of the ICF compartment is found within muscle cells. The primary
electrolytes of the ICF compartment are potassium and phosphate. The ICF
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compartment contains only small quantities of sodium and chloride ions and almost
no calcium ions. The cells contain 4 times as much protein as the plasma.
Intracellular fluid provides body cells their turgor as well as a medium within which
biochemical reactions can take place.

2.3 Extracellular fluid (ECF)


All the fluids outside the cells are collectively called the extracellular fluid. Together
these fluids account for about 20% of the body weight, or about 14 litres. The ECF
supports the cells and allows transport of nutrients and waste products. The
extracellular fluid is divided into the interstitial fluid and the blood plasma. There is
another small compartment of fluid that is referred to as transcellular fluid.

Interstitial fluid (IF)


It is the fluid that surrounds the cells in the various tissues of the body. It is about
three-quarter of the ECF volume. It includes the water contained within the bone and
dense connective tissue. The remaining one quarter of the ECF is fluid inside the
blood vessels, that is, plasma.

Transcellular fluid (TCF)


This is the fluid located in special compartments of the body. It is usually considered
to be a specialised type of extracellular fluid. Their total volume is about 0.3 litre and
they serve important functions. This compartment includes fluid in the synovial,
peritoneal, pericardial, and intraocular spaces, as well as the cerebrospinal fluid.

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2.4. Measurement of Body Fluid Compartment
This is usually done using dilution technique. The plasma volume, ECF volume and
the total body water (TBW) can also be measured using dilution technique. The
interstitial and intracellular fluid volumes can then be derived as follows:
ICF = TBW - ECF
IF = ECF - plasma volume
Plasma volume can be measured using dyes that bind to plasma protein e.g. Evans
blue (T1824). Plasma volume can also be measured by injecting serum albumin
labelled with radioactive iodine. Total body water can be measured using the dilution
technique involving a substance that will mix evenly with all the body fluid
compartments. Radioactive water (tritium, 3H 2 O) or heavy water (deuterium, 2H 2 O)
can be used for this measurement. Another substance that has been used to measure
total body water is antipyrine.

We can measure the ECF volume using any of the many substances that mix freely
in the plasma and interstitial fluid but do not readily pass through the cell membrane.
Such substances include radioactive sodium, radioactive chloride, and inulin. Also,
we can use mannitol and sucrose to measure ECF volume. However, we cannot
measure interstitial fluid volume directly even though it can be calculated as the
difference between the ECF volume and plasma volume.
Interstitial fluid volume = ECF - plasma volume
The ICF volume also cannot be measured directly but it can be determined as the
difference of TBW and ECF.
ICF volume = TBW- ECF volume.
Blood volume can be calculated if the hematocrit (the fraction of the total blood
volume composed of cells) is known, using the following equation:
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Total Blood Volume = Plasma Volume
1 - Hematocrit
For example, if plasma volume is 3liters and hematocrit is 0.40, total blood volume
would be calculated as:
3liters
1 - 0.4 = 5liters.
Another way we can measure blood volume is to inject into the circulation red blood
cells that have been labelled with radioactive material. After this has mixed in the
circulation, the radioactivity of a mixed blood sample can be measured, and the total
blood volume can be calculated using the dilution principle. A substance frequently
used to label the red blood cells is radioactive chromium (51Cr), which binds tightly
with the red blood cells.

2.5 Blood
Blood is the fluid that circulates through the heart, arteries, capillaries and veins,
carrying nutrients and oxygen to the body cells. It also carries waste products of
metabolism, the most important of which is carbon dioxide, from the tissues to the
organs where such waste products are expelled from the body. Blood is made up of
yellow liquid called plasma and the cellular component. The cellular component of
blood is made up of three types of cells: the red blood cells (erythrocytes), the white
blood cells (leucocytes) and the platelets (thrombocytes). The “cells” in the blood are
also called corpuscles.

Functions of blood
i. It transports oxygen from air in the lungs to the tissues and carbon dioxide
from the tissues to the lungs where it is expelled.
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ii. It transports food materials such as glucose, fatty acids, amino acids,
vitamins and electrolytes from the gastrointestinal tract to body tissues where
they are utilised for body building and energy production.

iii. It helps in the exchange of water, electrolytes and hydrogen ions between
the various body compartments.

iv. It also plays important role in the homeostasis of other body constituents
such as glucose, hormones etc.

v. It helps in transporting heat from one part of the body to the other.
vi. It transports humoral, antibodies, chemical agents, enzymes, and cellular
elements that are important in the defence of the body against infection or
invasion by non-infective foreign tissues of organisms.

Blood can be divided into two main parts namely cells or formed elements and
plasma. Cells consist of red corpuscles, white corpuscles, and platelets. Plasma is a
mixture of water, amino acids, proteins, carbohydrates, lipids, vitamins, hormones,
electrolytes, and cellular wastes.

Haemopoiesis/Hematopoiesis
This is the formation of blood cells. In the bone marrow are cells called Pluripotent
haemopoietic stem cells (PHSC) from which all circulating cells are derived. The
PHSC undergoes successive divisions to form different blood cells. Growth and
reproduction of the stem cell is controlled by multiple proteins called growth
inducers. e.g. Interleukin - 3 (IL - 3). Another set of proteins called the

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differentiation inducers causes one type of stem cell to differentiate towards a final
type of adult blood cell. e.g. erythropoietin.

Figure 2.2.2: Development of various formed elements of the blood from bone marrow cells

Red blood cells


Normal red blood cells (erythrocytes) are biconcave discs with a diameter of 7.2
microns and a thickness of 2.2 microns that contain one-third oxygen-carrying
haemoglobin by volume. When oxygen combines with haemoglobin, the resulting
oxyhaemoglobin is bright red. Red blood cells discard their nuclei during
development.
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The typical red blood cell count is 4,600,000 — 6,200,000 cells per mm3 for males
and 4,500,000 — 5,100,000 cells per mm3 for females. The number of red blood
cells is a measure of the blood's oxygen-carrying capacity. With age, red blood cells
become increasingly fragile and are damaged by passing through narrow capillaries.
Macrophages in the liver and spleen phagocytise damaged red blood cells.
Haemoglobin from the decomposed red blood cells is converted into heme and
globin. Heme is decomposed into iron and biliverdin. Iron is recycled into new
haemoglobin or stored in the liver. Some biliverdin is converted into bilirubin.
Biliverdin and bilirubin are excreted in bile as bile pigments.

Erythropoiesis
This is the process of production of red blood cells. In the embryo and foetus, red
blood cell production occurs in the yolk sac, liver, and spleen; in the adult it occurs
in the red bone marrow. The average life span of a red blood cell is 120 days. The
three cellular components of blood, that is red blood cells, white blood cells and
platelets, originate from the same primitive or pluripotential haemopoietic stem cells
(PHSC), from the bone marrow. The PHSC is uncommitted and can develop into
any of the three cell types. The PHSC however differentiates (through mitotic
divisions), to form committed stem cells. The committed stem cells at this stage
are called colony-forming unit (CFU). A committed stem cell that produces
erythrocytes is called a colony-forming unit- erythrocyte or CFU-E, while those that
produce granulocyte and monocyte are called CFU-GM, those for platelets
(megakaryocytes) are called CFU-M.

In erythropoiesis, large numbers of the first cell that can be identified as belonging to
the red blood series, the proerythroblast, are formed from the CFU-E stem cells
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under appropriate stimulation. Once the proerythroblast has been formed, it divides
several more times and goes through many stages of development before it
eventually forms many mature red blood cells. Thus, the proerythroblast goes
through the following stages:

Figure 2.2.3: Genesis of normal red blood cells (RBCs)

As the proerythroblast undergoes successive division, the daughter cell becomes


smaller than its precursor, and its nucleus occupies a smaller proportion of the
total cell volume. These tend towards smaller cells and nuclei with successive cell
types in the series, the early, intermediate and late erythroblast. The late erythroblast
loses its nucleus and thereby becomes a reticulocyte. The reticulocytes enter the blood
by squeezing between the endothelial cells (diapedesis) of the sinusoids in the bone
marrow. The cells lose their endoplasmic reticulum and mature into erythrocytes in one
or two days. From proerythroblast to erythrocyte takes three to four days. For the red
cell count to remain constant, the rate of production of red cells must equal the rate of
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destruction of old cells. The rate of production is indirectly controlled by the oxygen
content of the blood.

Factors promoting erythropoiesis


Factors important in promoting erythropoiesis are erythropoietin, vitamins, iron, other
hormones, proteins and certain trace elements.

Erythropoietin
This is a hormone that is also known as erythropoiesis stimulating hormone. It is the
main regulator of erythropoiesis in our body. It is a glycoprotein and has a molecular
weight of 34,000. It is formed mostly in the kidneys (90%) and the remaining 10% is
formed in tissues outside the kidneys, mainly in the liver. The main stimulus for the
production of erythropoietin is oxygen deficiency (reduced oxygen delivery) to the
tissues which can be caused by hypoxia, bleeding, anaemia etc. Erythropoietin then
stimulates the haemopoietic tissues to form more red blood cells. Erythropoietin also
causes the proerythroblasts to divide more rapidly and proceed to mature erythrocytes.
The total number of red blood cells remains relatively constant due to a negative
feedback mechanism utilising the hormone erythropoietin, which is released in
response to low oxygen levels detected in the kidneys and liver.

White blood cells


The leukocytes, also called white blood cells, are the mobile units of the body’s
protective system. White blood cells (leukocytes) help defend the body against
disease. Five types of white blood cells are in circulating blood and are distinguished
by size, granular appearance of the cytoplasm, shape of the nucleus, and staining
characteristics. The types of white blood cells are the granular neutrophils,
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eosinophils, and basophils, and the agranular monocytes and lymphocytes.
Normally a cubic milliliter of blood contains 5,000 to 10,000 white blood cells.

Formation of granulocytes
The granulocytes are formed in the red bone marrow. The entire maturation process
from myeloblast to neutrophil takes about 3 days. The sequence of cell
division/differentiation which gives rise to granulocytes is as follows:
Pluripotent haemopoietic stem cell

Unipotent committed precursor stem cell

Myeloblast

Promyelocyte

Myelocyte

Metamyelocyte

Neutrophil Eosinophil Basophil

Granulocyte and macrophage colony- stimulating factors.


The production of red and white blood cells is regulated with great precision in
healthy humans. The production of granulocytes is rapidly increased in infections by
bacteria, viruses, fungi or parasites. The proliferation and maturation of the cells that
enter the blood from the marrow are regulated by glycoprotein growth factors or
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hormones that cause cells in one or more of the committed cell lines to proliferate
and mature.
Three other factors, called colony-stimulating factors (CSFs) cause committed stem
cells to proliferate and mature. These are
(i) Granulocyte-macrophage CSF. (GM-CSF)
(ii) Granulocyte CSF (G-CSF)
(iii) Macrophage (M-CSF)
Interleukins also play important roles in haemopoiesis. Interleukins IL-1 and IL-6
followed by IL-3 act in sequence to convert pluripotent uncommitted stem cells to
committed progenitor cells. Interleukins IL-4 and IL-5 also play their roles.
Neutrophils have red-staining fine cytoplasmic granules and a multilobed nucleus;
they comprise 54-62% of leukocytes. It is a very motile and actively phagocytic cell
that ingests bacteria and other microorganisms, small particulate matter and fibrin.
The cytoplasmic granules of neutrophils appear to be the special types of lysosomes
containing hydrolytic enzymes which are used to destroy bacteria and other
microorganisms. Because of their motility, phagocytic activity and large numbers,
neutrophils play a key role in the body’s defence against bacterial invasion. The
neutrophils seek out, ingest and kill bacteria and have been called the body’s first line
of defence against bacterial infections. Neutrophils also release thromboxanes
that are vasoconstrictors and platelet- aggregating agents.

Eosinophils have coarse granules that stain deep red, a bilobed nucleus, and make up
only 1-3% of circulating leukocytes. They are weak phagocytes, and they exhibit
chemotaxis. They are less motile than the neutrophils. The cytoplasmic granules of
eosinophils contain many enzymes such as oxidases, peroxidases, and phosphatases,
indicating that the primary function of the eosinophil is detoxification of foreign
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proteins and other substances. Eosinophils are more abundant in connective tissues
than in the blood, particularly the lung, the mammary glands, omentum, and the inner
wall of the small intestine. Basophils have fewer granules that stain blue; they account
for less than 1% of leukocytes. The basophils in the circulating blood are similar to the
large tissue mast cells located immediately outside many of the capillaries in the body.

Both mast cells and basophils liberate heparin into the blood, a substance that can
prevent blood coagulation. The mast cells and basophils also release histamine, as well
as smaller quantities of bradykinin and serotonin. In allergic reactions, the antigen-
antibody reaction cause mast cells or basophils to rupture and release exceedingly large
quantities of histamine, bradykinin, serotonin, heparin, slow-reacting substance of
anaphylaxis and a number of lysosomal enzymes. These substances then cause local
vascular and tissue reactions that cause many, if not most, of the allergic
manifestations. Monocytes are the largest blood cells, have variably-shaped nuclei, and
make up 3-9% of circulating leukocytes. They migrate into tissues, enlarge up to five
times and develop numerous cytoplasmic granules (lysosomes). These cells are called
macrophages and they are much more powerful phagocytes than neutrophils.
Macrophages have a powerful lysosomal lipase which breaks down the lipid-rich cell
membranes of many bacteria. Many macrophages become fixed within tissues. These
stationary phagocytic cells are called tissue macrophage system.

Tissue macrophage are found virtually in all areas of the body especially in the skin
and subcutaneous tissue, the lymph nodes, alveoli of the lungs, the liver sinuses
(Kupffer cells), the spleen and marrow and in the brain. Lymphocytes are long-lived,
have a large, round nucleus, and account for 25-33% of circulating leukocytes. They
are of two main types:
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(1) B-lymphocytes, which are for humoral immunity i.e. they synthesise circulating
antibodies.
(2) T-lymphocytes, which are processed by or in some way dependent on the thymus
gland. They are responsible for cell-mediated immunity i.e. the production of
lymphocytes which are sensitised against specific antigens.

The life of the granulocytes after being released from the bone marrow is normally 4 to
8 hours circulating in the blood and another 4 to 5 days in tissues where they are
needed. The monocytes also have a short transit time, 10 to 20 hours in the blood,
before wandering through the capillary membranes into the tissues. Once in the tissues,
they swell too much larger sizes to become tissue macrophages, and, in this form, can
live for months unless destroyed while performing phagocytic functions. The
lymphocytes have life spans of weeks or months; this life span depends on the body’s
need for these cells. The platelets in the blood are replaced about once every 10 days;
in other words, about 30,000 platelets are formed each day for each microliter of
blood.

Platelets
Platelets are one of the formed elements of blood the others being red blood cells and
white blood cells. Platelets are formed from the megakaryocyte cells in the marrow
by pinching-off of bits of cytoplasm and extruding them into the circulation.
Platelets are not nucleated and they have a diameter of 2-3μ. The normal platelet
count is 200,000- 400,000 per mm3. Platelets contain glycogen, lysosomes and two
types of granules: dense granules and alpha-granules. The dense granules contain
ADP, serotonin and calcium. The alpha-granules contain clotting factors and other

102
proteins. Platelets have the ability to collect at the site of injury (platelet aggregation)
and discharge the contents of their granules (platelet release).

Functions of platelets
(i) They prevent blood loss by adhering to the vessels walls and forming aggregate
plugs.
(ii) On damage, they undergo the release reaction and release amines (histamine,
serotonin, and adrenaline), adenine nucleotides (ADP) and phospholipids.
(iii) They bring about clot retraction (platelet contractile protein).
(iv) They contribute to endothelial integrity
The production of platelet is regulated by thrombopoietin or thrombopoietic
stimulating factor (TSF) which is present in the blood. Thrombopoietin increases the
formation of megakaryocytes from committed stem cells in the bone marrow.

Plasma and Plasma proteins.


Plasma
Plasma is the fluid portion of the blood. It is straw-coloured. It is part of the
extracellular fluid, although it is found in the intravascular space. Its composition is
similar to that of the interstitial fluid, except for a much higher concentration of
proteins in the plasma.

Composition of plasma
Plasma is composed of
(i) Water
(ii) Electrolytes- Na+, K+, Cl-, HCO 3- , SO 4 , PO 4 , Ca++, Mg++
(iii) Plasma proteins- albumin, globulins and fibrinogen
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(iv) Products of digestion- glucose, free- fatty acids, amino-acids
(v) Hormones that have been released into the blood
(vi) Dissolved gases, especially carbon dioxide and a little quantity of oxygen
(vii) Metabolic waste products such as urea, uric acid, bilirubin.

Plasma proteins
The plasma proteins consist of albumin, globulin and fibrinogen. The total plasma
protein concentration is 64-83g per litre (about 6-8g per 100ml). The globulin
fraction can be subdivided into alpha 1 , alpha 2 , beta 1 , beta 2 , and gamma globulins.
The molecular weight of albumin is 69,000 while that of fibrinogen is 340,000.
Because of their large molecular size, the plasma proteins do not normally pass
through the capillary wall into the interstitial space. The proteins remain in the blood
vessels and exert an osmotic force of about 25 mmHg across the capillary wall
(oncotic pressure) that tends to pull the fluid from the interstitial space into the
intravascular space.

Functions of plasma proteins


(i) Exerts an osmotic pressure of 25 mmHg (oncotic pressure) that helps to pull
water from the tissue spaces back into the blood.
(ii) Helps to transport various substances, e.g. are albumin-transports calcium,
bilirubin, alpha-globulin-transports cortisol, thyroxine, Vitamin B 12 . beta-globulin
transports iron (transferrin) cholesterol, lipids, insulin, the fat soluble vitamins A, D,
and K.
(iii) Plasma proteins act as blood buffers. They are responsible for 15% of the
buffering capacity of the blood.
(iv) The proteins contribute to the viscosity of the blood.
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(v)The globulins are the antibodies that defend the body against foreign antigens.

Blood groups.
We humans can be divided into different groups based on the type of antigen on the
surface of our red blood cells and the antibodies in our plasma. There are two main
systems of blood group- the A.B.O. system and the Rhesus system.

The ABO system


This system is based on the presence of A, B, or A and B agglutinogens (antigens)
on the surface of the red blood cells. On the basis of these agglutinogens, we can be
classified into four groups: groups A, B, AB and O. The plasma of each group
contains agglutinin (antibody) that is opposite in name to the antigen on the surface
of its red blood cells. For example, group A contains A antigen on the surface of its
red blood cells and B antibody in its plasma. The distributions in the different groups
are summarised in the table below
Blood Group Agglutinogen Agglutinin
A A Anti - B
B B Anti - A
AB AB None
O None Anti A and Anti B
The blood group antibodies are gamma globulins, mostly of the IgM types and they
are produced by the same cells that produce antibodies to other antigens present or
introduced into the body. The A, B, O blood group is inherited in a Mendelian
fashion. The three genes involved are A, B, and O.

The six possible phenotypes with the corresponding blood groups are:
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Phenotype Blood Group
AA A
AB AB
BB B
AO A
OO O
BO B

Determination of blood group


We can determine blood group using anti-A and Anti-B sera. A drop of the anti-
serum is placed on a tile (or slide). The blood whose group is to be determined is
diluted 1 in 20 with saline (using a white cell pipette of a haemocytometer). A drop
of the diluted blood is added to the anti-sera and left for about 10 minutes. At the end
of 10 minutes, the mixture is stirred with a glass rod and it is noted whether
agglutination has taken place. The typical agglutination pattern is
shown in the table.

RBC of Unknown Group Anti- A Anti-B


Blood Group
RBC - - O
RBC + - ARBC
- + B
RBC + + AB
- means no agglutination
+ means agglutination is present

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RBC of
Unknown Anti- A Anti-B Blood Group
Group
RBC - - O
RBC + - A
RBC - + B
RBC + + AB

- means agglutination is absent


+ means agglutination is present

Rhesus blood group


In Rhesus blood group, there are six types of agglutinogens named C, D, E, and c, d,
e. The first three are dominant and the last three are recessive. Of the dominant
antigens, the main one is the D antigen. 85% of the population of all white people
have the D agglutinogens and are Rhesus positive. The remaining 15% don’t have D
agglutinogens and are Rhesus negative. In American blacks, the percentage of Rh
positives is about 95%, whereas in some African blacks, it is virtually 100%. Unlike
in the A, B, O system, the Rhesus blood group has no naturally occurring antibody in
the plasma to the D antigen. The antibody is only developed when a Rhesus negative
person is exposed to a Rhesus positive blood. Any of the A, B, O, blood groups can
be Rh positive or Rh negative.

Thus, we can have A + ve, A -ve, B +ve, B-ve etc. Rhesus blood group is of great
importance in women in relation to pregnancy. The problem is mainly in the Rhesus

107
negative women. If she has been previously sensitised (by Rh positive blood
transfusion or Rh positive baby where foetal and maternal blood mixed together at
parturition), if the next pregnancy is an Rh positive foetus, the foetus may develop
severe haemolysis and jaundice due to Rh incompatibility or be born dead (hydrops
foetalis). An Rh positive woman does not have such problem.

Blood transfusion
This is the process whereby one person gives blood to be passed into the body of
another person. The person who gives blood for someone else’s use is called “donor”
and the person to whom blood is given is called “recipient”. Blood transfusion may
be indicated in cases of excessive blood loss, severe anaemia or during surgical
operation. To ensure that there is no agglutination, it is desirable that a recipient
should be given blood of the same group as his own. Since this is not always
possible, a recipient can be given blood from another group provided the agglutinins
in the recipients’ plasma will not react with the agglutinogens on the donors cells.
For instance, a recipient who is group A cannot be given a group B blood since the
Anti-B in the group A plasma will react with the B agglutinogens on the group B
donor cells. Apart from people with the same blood group giving blood to
themselves, the other possibilities in transfusion are as
follows:
A

O AB

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From the above diagram, you will note that group O can give blood to all the other
groups, including group O, hence this group is called universal donor. Similarly,
group AB can receive blood from all other groups; hence the group is called
universal recipient. It is necessary to explain here that although blood group O
contains anti-A and anti-B, it can be given to blood groups A, B and AB with A
and/or B agglutinogens because the volume of the donor blood is far less than the
total blood volume of the recipient. Hence, the agglutinin titre in the donor blood is
considerably diluted by the larger volume of the recipient’s blood so that it is no
longer strong enough to cause agglutination.

Hazards of blood transfusion


Blood transfusion can be associated with some hazards, although, with great care,
such hazards are indeed uncommon. The possible hazards are:
(a) Incompatibility- this can be of the A B O; Rhesus; or expired blood types.
Inappropriate storage (excessive heat or cold) can also cause incompatibility.
(b) Overloading of the circulation- this occurs if large volume of blood is given too
rapidly
(c) Air embolism
(d)Transmission of infection e.g. malaria, AIDS virus.
(e) Allergic reaction
(f) Sensitisation, in case of Rh-positive blood to a Rh-negative person on the first
occasion.
(g) Febrile reaction due to pyrogens. This can be due to contamination of the blood
giving set

109
ITQ 1: What is blood transfusion?
ITA 1: This is the process whereby one person gives blood to be passed into the body of another
person.

3.0 Conclusion/Summary
Body fluids constitute about 2/3 of the body weight and are distributed in different
compartments of the body. They majorly consist of the intracellular and extracellular
fluids which can be measured in different ways. ABO blood group system is the
major blood group system that has clinical significance in blood transfusion.

4.0 Self-Assessment Questions


1. What is erythropoiesis ?
2. Describe the Rhesus blood group? Discuss the different types of body fluid
compartments
3. Briefly describe the measurement of body fluid compartments
4. Discuss the ABO blood group system

5.0 References/Further Readings


Sembulingam K., and Prema S. (2012), Essentials of Medical Physiology. 6th Ed.
India: Jaypee Brothers Medical Publishers.
Kim E. et.al (2010), Ganong’s Review of Medical Physiology 23rd Ed. United States:
The McGraw-Hill Companies, Inc
Guyton A. and John E. (2006) Textbook of Medical Physiology 11th Ed. Philadelphia:
Elsevier Inc.

110
STUDY SESSION 3
Haemostasis
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1- Haemostasis
2.2- Formation of Platelet plug
2.3- Formation of blood clot and clot retraction
2.4- Repair of blood vessel endothelium
3.0 Conclusion/Summary
4.0 Self-Assessment Questions
5.0 Additional Activities (Videos, Animations & out of Class activities)
6.0 References/Further Readings

Introduction:
The body is prone to injury from daily activities and there must be a way of
regulating blood loss otherwise the body will lose the source of life. In this Session,
we are going to learn more about the process of how the body controls loss of blood.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Explain the concept of Haemostasis
2. Describe the sequence of events that cumulate in haemostasis.
3. Explain the process of Vascular spasm
4. Describe the formation of platelet plug
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5. Discuss the formation of blood clot and clot retraction
6. Describe the repair process of blood vessel endothelium

2.0 Main Content


2.1 Haemostasis
This is the process by which blood loss from our body is prevented when a blood
vessel is cut or Ruptured. Following injury to a blood vessel, the sequence of events
in haemostasis is as follows:
i. Vascular spasm
ii. Formation of platelet plug
iii. Formation of blood clot
iv. Growth of fibrous tissue into the clot to form a permanent seal at the point of
vessel damage
v. Removal of excess fibrous tissue (fibrinolysis)
vi. Repair of blood vessel endothelium
ITQ 1: What is haemostasis?
ITA 1: This is the process by which blood loss from the body is prevented when a blood vessel is
cut or ruptured.

2.2 Vascular spasm


This is a process in which the injured vessel contracts to narrow or obliterate its
lumen, so as to reduce or stop blood loss altogether. Right after a blood vessel is
injured, it undergoes a powerful vasoconstriction (vessel narrowing). Because the
vasoconstriction is so strong and sudden, it is often called a vascular spasm. This
spasm and narrowing of the vessel lumen immediately slows the rate of blood loss,
because less blood is now flowing through the constricted, injured vessel.

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This spasm lasts about 20 to 30 minutes during which time, the next two processes
formation of platelet plug and formation of blood clot would have occurred thereby
providing a more effective seal for the damaged vessel.

2.3 Formation of Platelet Plug


There are about 200,000 to 400,000 platelets per mm3 of blood. When a vessel is cut
open, the collagen fibres in its wall are torn. Platelets in the surrounding blood
become sticky and attach to the shredded ends of the collagen fibres. As they pile up,
a platelet plug is created that partially closes off the hole in the vessel wall. Platelet
also releases ADP and enzyme that cause formation of thromboxane A.
Thromboxane A and ADP further cause the sticking together of platelet leading to
platelet plug.

2.4 Formation of Blood Clot and Clot Retraction


This involves coagulation of the blood that comes out of the blood vessel at the site
of injury. Injury to the vessel wall creates a chemical called prothrombin activator.
As its name indicates, prothrombin activator converts prothrombin (an inactive
clotting enzyme) into an active clotting enzyme called thrombin. Thrombin acts
upon fibrinogen, which are soluble or dissolvable within the bloodstream to form
fibrin. Fibrin filaments are insoluble (not dissolvable) within the blood plasma.
Hence, the fibrin filaments settle out of the blood and are deposited as a fibrin
meshwork over the platelet plug. More circulating platelets, as well as a few red
blood cells (RBCs), get stuck in this fibrin meshwork.

The process by which our body prevents blood loss occurs through two mechanisms
to produce a definite fibrin clot. Disorders in either system can cause diseases that
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cause either too much or too little clotting. They are the Intrinsic and Extrinsic
mechanisms. They involve the interplay and activation of proteins or clotting factors.
They are;
Factor I= Fibrinogen
Factor II= Prothrombin
Factor III= Tissue factor
Factor IV= Calcium
Factor V = Labile factor
Factor VI - Does not exist as it was named initially but later on discovered not to
play a part in blood coagulation.
Factor VII = Stable factor
Factor VIII = Antihemophilic factor A
Factor IX = Antihemophilic factor B or Christmas factor (named after the first
patient in whom the factor deficiency was documented)
Factor X = Stuart Power factor
Factor XI = Antihemophilic factor C
Factor XII = Hageman factor
Factor XIII = Fibrin stabilising factor

Extrinsic Pathway
In the Extrinsic pathway, tissue trauma leads to release of two factors— tissue
factors (TF), a proteolytic enzyme and tissue phospholipids. The latter is mainly
from damaged cell membranes. Factor X, in the presence of tissue factors, factor VII
and tissue phospholipids is converted to activated factor X. Activated factor X reacts
with tissue phospholipids and Factor V leading to formation of thromboplastin
(prothrombin activator).
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(i) Tissue damage TF + TP
(ii) Factor X TF, VII and TP Xa
(iii) Xa + TP + V Prothrombin activator
Where TF = Tissue factor
TP = Tissue phospholipids VII= Factor VII
V = Factor V
Xa = Activated Factor X

Intrinsic Pathway
In the Intrinsic pathway,
(a) Contact of blood with collagen or a foreign surface converts Factor XII to
activated factor XII, a proteolytic enzyme. Also, trauma leads to platelets
aggregation and release of platelet phospholipid or platelet factor.
(b) Next, Factor XI is acted upon by activated Factor XII to form Activated Factor
XI.
(c) Factor IX is acted upon by activated Factor XI and converted to activated Factor
IX.
(d) Factor X is acted upon by activated Factor IX, Factor VII and platelet
phospholipids and converted to Activated Factor X.
(e) Activated Factor X reacts with Factor V and platelet phospholipids to
form thromboplastin (prothrombin activator).

Conversion of prothrombin to thrombin


Prothrombin is formed in the liver and vitamin K is required in its formation.
Prothrombin is converted by prothrombin activator (formed in the extrinsic and

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intrinsic pathways) in the presence of calcium ions to thrombin. Platelets also play
an important role in the conversion of prothrombin to thrombin.

Conversion of fibrinogen to fibrin


Fibrinogen is also formed in the liver. Thrombin, a proteolytic enzyme, acts on
fibrinogen and converts it to fibrin monomer. The fibrin monomers join one another
to form long fibrin threads. These fibrin threads form a mesh-work that traps blood
cells, platelets and plasma. The initial fibrin thread can be easily broken, but it is
acted upon by fibrin stabilising factor (factor XII). The factor XII is first activated by
thrombin and the activated factor XII act on the fibrin threads to strengthen them so
that they can no longer be broken easily.

Clot retraction
This is the final step in the clotting process. The clot formed above is soft and jelly-
like. The fibrin threads in the clot contract a few minutes after the clot is formed and
squeeze out most of the fluid (serum) so that what is left is a firm clot. Platelets are
important in clot retraction and low platelets count can lead to failure of clot
retraction. Finally, excess fibrin that may be occluding part of the blood vessel
lumen is removed (fibrinolysis) and the damaged endothelium is replaced by a new
one.

Growth of fibrous tissue into the clot


The clot is soon invaded by fibroblasts leading to formation of dense fibrous tissue.
The latter completes the final sealing of the cut in the blood vessel.

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Fibrinolysis
If a small blood vessels is involved, the fibrous tissue may permanently occlude the
entire lumen of the vessel, but if a big blood vessel is involved, the fibrous tissue at
the site of vascular injury is retained, but excess fibrous tissue in clots that must have
extended to the lumen of the vessel is dissolved by the enzyme plasmin, so that free
flow of blood can occur once more.

2.5. Repair of Blood Vessel Endothelium


This is the final step in the process of haemostasis. The damaged endothelium is
replaced by the formation of a new lining.

ITQ 2: What is vascular spasm?


ITA 2: This is a process in which the injured vessel contracts to narrow or obliterate it lumen, so
as to reduce or stop blood loss altogether.

3.0 Conclusion/Summary
In this session, we have studied how blood loss is controlled by several mechanical
and chemical mechanisms. You have learnt that blood loss is controlled through
vascular spasm, formation of blot clot, growth of fibrous tissue into the clot to
form a permanent seal at the point of vessel damage, fibrinolysis and repair of
blood vessel endothelium.

4.0 Self-Assessment Questions


1. List the sequence of events in haemostasis?
2. Briefly describe fibrinolysis
3. Describe the intrinsic pathway
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4. Outline the sequence of events that occur to achieve repair of an
injured blood vessel.

7.0 References/Further Readings


Sembulingam K., and Prema S. (2012), Essentials of Medical Physiology. 6th Ed.
India: Jaypee Brothers Medical Publishers.
Kim E. et.al (2010), Ganong’s Review of Medical Physiology 23rd Ed. United States:
The McGraw-Hill Companies, Inc
Guyton A. and John E. (2006) Textbook of Medical Physiology 11th Ed. Philadelphia:
Elsevier Inc.

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STUDY SESSION 4
Immune System
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1- Immunity
2.2- Active immunity
2.3- Mechanism of action of antibodies
2.4- Passive immunity3.0 Tutor Marked Assignments (Individual or Group)
3.0 Conclusion/Summary
5.0 Self-Assessment Questions
6.0 Additional Activities (Videos, Animations & out of Class activities)
7.0 References/Further Readings

Introduction:
You may have wondered how your body is able to survive in an environment with
microorganisms and the toxins of these organisms that are often dangerous to the
survival of the body. It is your body’s immune system helps to fight microorganisms
and develop some resistance to them. In this unit, you are going to learn more about
the different types of immunity and the mechanism of antibody function.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Explain the concept of Immunity
2. Discuss different types of Immunity
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3. Explain the difference between Passive immunity and Active immunity
4. Discuss the Mechanism of action of antibodies

2.0 Main Content


2.1 Immunity
This is the ability of our body to resist almost all types of organisms or toxic
substances that tend to damage our tissues or organs. It is classified into 2 major
groups: (a) Active immunity and (b) Passive immunity.

ITQ 1: What is immunity?


ITA 1: This is the ability of our body to resist almost all type of organisms or toxic substances that
tend to damage the tissue or organs.

2.2 Active Immunity


This is further divided into innate and acquired immunity.

2.2.1 The innate immunity


This involves all the processes already present in the body of an organism that are
directed at protecting the body of the organism, e.g. (i) the resistance of the skin to
invasion of organism (ii) destruction in the stomach of foreign organism entering the
gut through the mouth by acid secretion (iii) phagocytosis of bacteria and other
foreign invaders by white cells or cells of the tissue macrophage system (iv) also
present in the blood are certain chemicals such as lysosomes that attack foreign
organisms or toxins.

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2.2.2 Acquired immunity
This type of immunity is not present naturally in the body. It is activated when the
body is exposed to foreign organism. There are two types of acquired immunity
namely cellular and humoral immunity. It is the product of the body lymphocyte
system.

2.2.3 Cellular immunity


It plays an important role in the body’s defence against viral, bacteria, fungi
infection and in transplant rejection. T-lymphocyte function may be suppressed by
steroid hormones or by the drug azathioprine. Such drugs are used to prevent
rejection of grafted tissue or organelles. Cellular immunity is mediated by T-
lymphocytes. T- lymphocytes derived from the stem cell in bone marrow migrate
to the thymus where it is processed and become immunologically competent.
During this process, each lymphocyte T develop specific reactivity against one
antigen, thus at the end of the process there are different T-lymphocytes with
specific reactivity against millions of different antigen. These processed
lymphocytes leave the thymus and spread throughout the body lodging in the
lymphoid tissue all over the body. These T-cells have thousands of receptor
molecules on its surface and antigens bind with these receptors. T-cells processed in
the thymus are screened to ensure that when released into circulation they do not
react with the body tissue. Those that fail the test are phagocytised instead of being
released.
There are four major types of T-lymphocyte
(1) Helper T cells
(2) Cytotoxic or killer T cells
(3) Suppressor T cells
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(4) Memory T cells

2.2.4 Helper T cell


They are the most numerous of the T cells and they constitute 70% of T-lymphocyte.
They serve as the main regulator of all immune functions. They do this by forming
series of products mediator called lymphokines or interleukins that act on the cell of
the immune system. The important interleukin secreted by the Helper T cell are IL-2,
IL-3, IL-4, IL-5, IL-6, granulocyte-monocyte cloning stimulating factor and the
interferon. When these lymphokines are absent, the remainder of the immune system
is paralysed. Helper T cells stimulate the growth of cytotoxic T cell and suppressor T
cell. This is mediated by the interleukin -2 with IL-4 and IL-5 playing subsiding role.
T cells also stimulate the B-cell growth and differentiation to form plasma cell and
antibodies. IL-4, IL-5, IL-6 are responsible for this action. The lymphokines of the
Helper T cells also ensure that macrophages slow down or stop their migration when
they reach the site of traction. They also activate the macrophages leading to
increase and infective phagocytosis. The cellular T cells are mediated by the
lymphocytes.

2.2.5 Cytotoxic cells


They are direct attack cell capable of killing microorganism. Each cell has antigen
specific surface receptors that bind tightly to organism or cell that contain their
specific binding antigen. They secrete hole-forming protein called perforins that
punch large holes in the membrane of the attached cell leading to inflow of fluid into
the cell and lysis. A single cytotoxic cell can kill hundreds of organism before its
own death. Cytotoxic T cell play an important part in destroying cancer cell, heart
transplant and other type of cells that are foreign to the body.
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2.2.6 Suppressor T cells
They suppress the function of both cytotoxic and helper T cells. They regulate the
activity of other cells preventing them from causing excessive immune reaction
which may damage the body tissue. The process by which suppressing T cell limit
the ability of immune to attack our own tissue is known as immune tolerance.

2.2.7 Memory T (or B) cells


These are cells that have been exposed to an antigen and are readily converted to
effector cells by later encounter with the same antigen.

2.2.8 Humoral immunity


This is mediated by B-lymphocytes. They are pre-processed in the liver during the
mid foetal life and in the bone marrow in the late foetal life and after birth. After pre-
processing, they migrate into the lymphoid tissue throughout the body. These cells
differentiate into plasma cell and memory B cells. The mature plasma cells are
responsible for the production of antibodies. These antibodies produced are called
immunoglobulins and there are five classes of immunoglobulins: IgM, IgG, IgA,
IgD, and IgE. IgG is the most abundant immunoglobulin in we humans (about 75%
of the total). Most antitoxins and virus antibodies belong to this class of
immunoglobulins. IgM are efficient in reacting with bacteria and foreign cells. IgA
protect the mucus membrane. IgE is involved in hypersensitivity such as asthma and
hay fever.

When an antigen is introduced into the body, antibodies appear in the blood after a
few days. The antibody produced on this first contact with antigen increases rapidly
to a peak which is not very high and then declines. If the same animal is injected
123
with the same antigen, the response occurs sooner and the amount of antibody
produced is much greater than the first exposure. This is the secondary immune
response. The more rapid appearance of antibody and the greater production are due
to the presence of long-lived B-lymphocytes called memory B cells.

2.3 Mechanism of Action of Antibodies


Antibodies act in two different ways: (i) by direct attack on the invader (ii) action of
the complement system that destroys the invader. The direct action of antibodies is
carried out in several ways which include: (i) agglutination (ii) precipitation (iii)
neutralisation and lysis. These direct actions of antibodies attacking antigenic
invader under normal condition are not strong enough to play a major role in
protecting the body against the invader. Most of the protections form the amplifying
effect of the complement system for antibodies action.

2.3.1 Complement system cell


Complement is a collective term of describing a system of 20 different proteins
many of which are enzyme precursors. The principal actors of the system are 11
proteins designated C1-C9 , B and D, shown in the figure below. The enzyme
precursors are normally inactive, but they can be activated in two ways: (i) the
classical pathway (ii) alternate pathway. The classical pathway is mediated when an
antigen-antibody reaction occurs. The activated C1
activates C2, C3, C4, and so on, setting in motion a “cascade” of reactions and the
enzyme produced cause the following effects:
(1) Phagocytosis
(2) Lysis
(3) Agglutination
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(4) Neutralisation
(5) Chemotaxis
(6) Activation of mast cells and basophils
(7) Inflammatory effects

Figure 2.4.1: Cascade of reactions during activation of the classical pathway of the complement
system

In the alternate pathway, antigen-antibody complexes are not required to activate the
system. This occurs in response to large polysaccharide molecules of some invading
organisms. These substances react with complement B and D forming an activated
product that activates factor C3 setting off the remainder of complement pathway.
This leads to the release of the same enzymes and the same effects as in the classical
pathway. Since the alternate pathway does not involve an antigen-antibody reaction,
it can function even before you are immunised against a particular organism.

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2.4 Passive Immunity
This can be achieved by injection of preformed antibody obtain from the plasma of
another blood cell or of an animal that have already been actively immunised against
a particular disease. Such antibody can be given in treatment of tetanus. Passive
immunity can also be transferred from a mother to a new born via colostrum.

ITQ 2: What are memories T-cells?


ITA 2: These are cells that have been exposed to an antigen and are readily converted to effector
cells by later encounter with the same antigen

3.0 Conclusion/Summary
In this lesson, we have studied the ways our body is able to resist infections through
active (innate or acquired) and passive immunity.

4.0 Self-Assessment Questions


1. What is innate immunity?
2. Explain humoral immunity?
3. Write an essay on different types of Immunity
4. In tabular form, differentiate between Passive immunity and Active immunity
5. Discuss the mechanism of action antibodies

5.0 References/Further Readings


Sembulingam K., and Prema S. (2012), Essentials of Medical Physiology. 6th Ed.
India: Jaypee Brothers Medical Publishers.

126
Kim E. et.al (2010), Ganong’s Review of Medical Physiology 23rd Ed. United States:
The McGraw-Hill Companies, Inc
Guyton A. and John E. (2006) Textbook of Medical Physiology 11th Ed. Philadelphia:
Elsevier Inc.

127
MODULE 3
Physiology of the Nervous System I
Contents:
Study Session 1: Physiology of the Central Nervous System: Brain
Study Session 2: The Spinal Cord, Cranial and Spinal Nerves
Study Session 3: Neurons and Neuronal Transmission
Study Session 4 & 5: The Autonomic Nervous System

STUDY SESSION 1
Physiology of the Central Nervous System: Brain
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Structural Organisation of the Brain
2.2 Functions of the Cerebrum
2.3 The Diencephalons
2.4 Emotion and Motivation
2.5 Memory
2.6 The Midbrain
2.7 The Hindbrain
2.8-Reticular Formation
3.0 Conclusion/Summary
4.0 Self-Assessment Questions and Answers
5.0 References/Further Readings

Introduction:
The Nervous system is one of the two major control systems of our body
in addition to the endocrine system. The central nervous system (CNS) is
128
composed of the brain and spinal cord. These receive input about the
external and internal environment from afferent neurons. The CNS sorts,
processes and transmits this information to the efferent neurons which
carry the instruction to glands or muscles to bring about the desired
response. The nervous system acts by means of its electrical signals to control the rapid
responses of the body. The brain, the first part of the central nervous system is
arranged in regions and subdivisions. In this unit, we shall be examining the functions
of the various regions of the brain and some higher brain functions.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Describe the structural organization of the brain
2. Explain the anatomical divisions of the cerebrum and their functions
3. Explain the lateralization and language function of the cerebral cortex
4. Explain the functions of the various parts of the encephalon
5. Explain the neural basis for the control of emotions, motivation and
memory
6. Describe the function of the midbrain
7. Identify the structures of the hindbrain and explain their functions
8. Explain the function of the Reticular formation.

2.0 Main Content


2.1. Structural Organization of the Brain
The central nervous system (CNS) consists of the brain and spinal cord. They receive
input from sensory neurons, and direct the activity of motor neurons. Association
neurons are present to "associate" appropriate motor responses with sensory stimuli. The
early embryo contains an embryonic tissue layer known as ectoderm
on its surface that will eventually form the epidermis of the skin and the
nervous system. As development continues, a groove appears in the

129
ectoderm along the dorsal midline of the embryo's body. The groove
deepens and by the twelfth day after conception has fused to form a neural
tube. The part of the ectoderm where the fusion occurs becomes a
structure separate from the neural tube and is called the neural crest.

The neural tube becomes the CNS later while the neural crest eventually
becomes the ganglions of the peripheral nervous system and other
structures. By the middle of the 4th week three distinct swellings are evident
on the anterior end of the neural tube which will form the forebrain,
midbrain and hindbrain. In the 5th week those three areas are modified to
form five regions. The forebrain divides into the telencephalon and
diencephalon. The midbrain is unchanged and the hindbrain divides into
the metencephalon and myelencephalon. These regions later become
greatly modified but the terms described are still used to indicate the
general regions of the brain.

Figure 3.1.1: The Human Brain

The telencephalon grows disproportionately in humans forming the two


enormous hemispheres of the cerebrum that cover the diencephalon, the

130
mid brain and some portion of the hindbrain. The CNS that began as a
hollow tube remains hollow even as the regions are formed. The hollow
parts divide into cavities called ventricles in the brain. These become
filled with cerebrospinal fluid. These cavities are connected to themselves
and are continuous with the central canal of the spinal cord. The central nervous system is
composed of grey and white matter. Grey matter is composed of high concentration of
neuronal cell-bodies and dendrites that do not have myelin sheath. Grey matter is
found in the surface layer (cortex) of the brain and deeper within the brain in neuronal
aggregations called nuclei. White matter consists of high concentration of
axon tracts. Because axons are usually myelinated they acquire the white
colour of myelin sheath. White matter lies under the cortex and also surrounds the nuclei.

As adults, our brain contains about 100 billion neurons, weighs about 1.5kg
and receives about 20% of the total blood flow to our body per minute.
The human brain is mostly water (75%) and has the consistency of gelatin.
It needs support for its existence. The brain is therefore protected by the
scalp, hair etc and by the reinforced bony cranium which is one of the
strongest structures in the body. It also floats shock-proof in cerebrospinal fluid and is
encased in three layers of cranial meninges — the dura mater, arachnoid mater and pia
mater.
ITQ 1:
1. What are the components of the CNS
2. Enumerate the Land marks of CNS
ITA 1:
1. The brain and spinal cord
2. Cerebrum, Mid brain, Hypothalamus, pons, thalamus, Medulla, Spinal cord and Cerebellum

2.2. Functions of the Cerebrum


The cerebrum is the only structure of the telencephalon, and the largest of
the brain regions (accounting for about 80% of the brain mass). It is
believed to be the centre for higher brain functions and the most
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sophisticated region of the brain. The cerebrum is divided into two halves,
the right and left cerebral hemisphere, and these two hemispheres are
connected to each other by a thick band of neuronal axons called corpus
callosum. The outer layer of the cerebrum is the cerebral cortex which
caps an inner core of white matter that houses the basal nuclei.

2.2.1. The Cerebral Cortex


The cerebral cortex is composed of a thin (2 to 4mm thick) outer covering of grey matter
overlying a thick central core of white matter. The cortex characteristically contains
numerous folds and grooves called convolutions. The elevated folds are called
gyri (singular-gyrus-) and the depressed grooves are called sulci (singular sulcus). Each
cerebral hemisphere is divided by deep sulci or fissures into lobes— 4 major ones and
probably an additional minor fifth lobe. The lobes are the frontal, parietal, temporal and
occipital lobes, which are visible from the surface and deep and fifth insula lobe.

The frontal lobe is the anterior portion of each cerebral hemisphere. It is


separated by a deep fissure called the central sulcus from the parietal lobe.
The precentral gyrus is located in the frontal lobe just in front of the
central sulus. This gyrus is responsible for voluntary motor control. Other
functions of the frontal lobe are that it is also responsible for speaking
ability and for elaboration of thought. The primary motor cortex in the
precentral gyrus on each side of the brain primarily controls muscles on
the opposite side of the body.

The parietal lobes situated directly on the top of the head behind the
frontal lobe and separated from the frontal lobe by the deep central sulcus.
The parietal lobes are primarily responsible for receiving and processing
sensory input such as touch, pressure, heat, cold and pain from the surface
of the body.

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The post-central gyrus, directly behind the central sulcus in the parietal
lobe is the primary area of the cortex responsible for the perception of
these sensations collectively called somaesthetic sensations. (i.e. those arising from
cutaneous, muscle, tendon, and joint receptors).

The temporal lobes are located at the sides of the head and contain the
auditory centres that receive sensory fibres from the cochlea of the ear.
The lobe is also involved in the interpretation and association of auditory
and visual information. The occipital lobes are located at the back of the
head and they are primarily responsible for vision and coordination of eye
movements.

The insula lobe is small and buried deep in the central sulucs. It is supposed to be
associated with memory, and visceral activities.

2.2.2 The Basal Nuclei


The basal nuclei is also known as basal ganglia. They are masses of grey matter
composed of neuron cell bodies located deep within the white matter of the
cerebrum. A nucleus (plural nuclei) refers to a functional aggregation of neuronal cell
bodies. The most prominent part of the basal nuclei is the corpus stratum. The major
function of the basal nuclei is a complex role in the control of
voluntary movement in the following ways :- (i) inhibiting muscle tone
throughout the body (to balance excitatory inputs); (ii) selecting and
maintaining purposeful motor activity while suppressing useless or
unwanted patterns of movement, and (iii) helping to monitor and coordinate slow
sustained contractions, especially motor neurons but they modify ongoing activity in
motor pathways.

133
The importance of the basal nuclei in motor control is evident in diseases involving this
region; the most common of which is Parkinson's disease. In this condition there is
deficiency of dopamine, a neurotransmitter in the basal nuclei. This lack makes the
basal nuclei unable to perform normal roles therefore the characteristic features of
Parkinson's disease manifest. These are (1) increased muscle tone or rigidity (2)
involuntary, useless or unwanted movements like resting tremors and (3) slowness in
initiating and carrying out motor behaviours.

2.2.3 Lateralisation and Kabgyage Language Function of the Cerebral Cortex


Each cerebral hemisphere controls the movements of the opposite side of the body
through motor fibres that originate in the precentral gyrus. In the same way, somaesthetic
sensations from each side of the body projects to the opposite side on the post central
gyrus due to crossing over (decussation of fibres). However, the two
hemispheres can receive information from both sides of the body because
they communicate with each other via the corpus callosum.

The cortical areas described so far appear to be equally distributed.


However, studies seen to indicate that a task can be successfully
performed by one side but not by the other. For example, the left
hemisphere has been shown to be the one in which most of the language
and analytical abilities reside as well as handedness. Fine motor control
seems to be more under the control of the left hemisphere. This is
evidenced by the fact that most of us are right handed and the left
hemisphere controls the right side of the body. It is these findings that led
to the concept of cerebral dominance. That is, since these various obvious
activities are controlled by the left hemisphere, then the left hemisphere
must be the dominant one.

However, further studies have shown that the right hemisphere is also specialised along
different less obvious lines. Therefore, rather than one hemisphere being dominant and
134
the other being subordinate, the two hemispheres appear to have complimentary functions.
The term cerebral lateralisation, or specialisation of function in one
hemisphere or the other, may be preferred, more recently to the term
cerebral dominance. Unlike the sensory and motor regions of the cortex, which are
present in both hemispheres, the areas of the brain responsible for language ability are
found in the left hemisphere in most of us.

Language is a complex form of communication in which written or spoken words


symbolise objects and convey ideas. It involves the integration of two distinct
capabilities namely, expression and comprehension, each of which is related to a
specific area of the cortex. The two areas are the Broca’s area and Wernicke's area.
Broca's area is responsible for speaking ability and articulation of speech. Broca's aphasia
is the result of damage to the Broca's area. Common symptoms include weakness of
the right arm and right side of the face. People with Broca's aphasia are reluctant to
speak, and when they speak, their speech is slow and poorly articulated. Speech
comprehension is not affected. They can understand a sentence but have difficulty
repeating it.
The Wernicke's area is located in the parietal lobe almost at the junction of
the parietal, occipital and temporal lobes. It is concerned with language
comprehension. It plays a critical role in understanding both spoken and
written messages. It is responsible for formulating coherent patterns of
speech that are transferred via a bundle of fibres to the Broca's area for
articulation. Wernicke's aphasia results in speech that is rapid but without
meaning. People with Wenicke's aphasia produce what has been
described as "word salad” e.g. real words chaotically mixed together or
made up words. Language comprehension whether written or spoken is
destroyed in Wernike's aphasia.

135
2.3 Diencephalon
Together with the telencephalon (cerebrum) the diencephalon constitutes
the forebrain. The diencephalon is almost completely surrounded by the
cerebral hemispheres and contains a cavity called the third ventricle. It
can be said to be the deep part of the cerebrum connecting the midbrain
with the cerebral hemispheres. It is composed of the thalamus,
epithalamus and hypothalamus. The pituitary gland is connected to the
hypothalamus.

The thalamus is composed of two egg-shaped masses of grey matter covered by a


thin layer of white matter. It is located in the centre of the cranial cavity directly
beneath the cerebrum and above the hypothalamus. It forms the lateral walls of the third
ventricle. The thalamus acts as an intermediate relay point and processing centre for
all sensory impulses (except smell) on their way to the cerebral cortex
from the spinal cord, brain stem, cerebrum, basal ganglia and other
sources. It screens out insignificant sensory impulses to appropriate areas
of the somato sensory cortex as well as to other regions of the brain. The
thalamus working with the brain stem and cortical association areas is
important in our ability to direct attention to stimuli of interest and so
forth.

The epithalamus is the dorsal portion of the diencephalon that contains a choroidplexus
over the third ventricle where cerebrospinal fluid is formed. It also contains the
pineal gland or epiphysis.

The hypothalamus is a small portion of the diencephalon located below


the thalamus, where it forms the floor and part of the central walls of the
third ventricle. It is a collection of specific nuclei and associated fibres.

This extremely important brain region is an integrating centre for many


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important homeostatic functions and serves as an important link between
the autonomic nervous system and the endocrine system. Specifically the
hypothalamus (1) controls body temperature; (2) controls thirst and urine
output; (3) controls hunger and food intake; (4) controls anterior pituitary
hormone secretion (5) produces posterior pituitary hormones (6) controls
uterine contraction and milk ejection; (7) serves as a major autonomic
nervous system coordinating centre which in turn affects all smooth
muscle, cardiac muscle and exocrine glands; and (8) plays a role in
emotional and behavioural patterns. In addition, centres in the
hypothalamus, contribute to the regulation of sleep, wakefulness, sexual
arousal and performance, and emotions as such anger, fear, pain and
pleasure.

The pituitary gland is located immediately inferior to the hypothalamus. It


is said to be derived embryonically from a down growth of the
diencephalon by means of a stalk neurons in the supraoptic and
paraveneticular nuclei of the hypothalamus produce two hormones— antidiuretic
hormone (ADH) and oxytocin. These hormones are transported to the
neurohypohypsis and stored and secreted in response to hypothalamic stimulation.

2.4 Emotion and Motivation


The parts of the brain that appear to be of paramount importance in emotional
states are the hypothalamus and the limbic system. The limbic system is not a separate
structure but refers to a ring of nuclei and fibre tracts that surround the brain stem and
are interconnected by intricate neuronal pathways. The structures of the limbic system
include the cingulate gyrus (part of cerebral cortex), amygdala, hippocampus and
septal nuclei. This complex interacting network of structures is associated
with emotions, basic survival and socio-sexual behavioural patterns motivation and
learning. It is the centre for basic emotional drives.

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There are few synaptic connections between the cerebral cortex and the structures of
the limbic system which may help explain the fact that we have so little control over
our emotions. However there is a closed circuit of information between the limbic
system and the thalamus and hypothalamus and they cooperate in the neural basis
of emotional states. Studies suggest that the hypothalamus and the limbic system are
involved in the following feelings and behaviours:
i. Aggression - stimulation of areas of the amygdala and particular areas of the
hypothalamus can both produce rage and aggression.
ii. Fear - electrical stimulation of the amygdala and hypothalamus produce fear.
Removal of the limbic system can result in absence of fear.
iii. Feeding - as discussed under hypothalamus.
iv. Sex - the hypothalamus and limbic system are involved in the regulation of
the sexual drive and behaviour.
v. Goal directed behaviours.
The concept of emotion encompasses subjective emotional feelings and moods
(anger, fear and happiness) plus the overt physical responses that
occur in association with these feelings. Such responses include specific
behavioural patterns (e.g. preparation for attack or defence) and observable
emotional responses (e.g. laughing, crying and blushing).

Motivation is the ability to direct behaviour towards specific goals that are aimed at
satisfying specific identifiable needs related to homeostasis. However most human
behaviours are shaped in a complex framework of unique personal gratifications blended
with cultural expectations.

2.5 Memory
Memory is the storage of acquired knowledge for later retrieval. The
neural change responsible for retention or storage is known as neural trace.
Concepts and not necessarily verbatim words are stored and so is recall.
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Storage of acquired information is believed to occur in at least two stages:-
short-term memory (STM) and long term memory (LTM). Short Term
Memory lasts for seconds to hours. Long term memory is retained for
days to years. When knowledge is first acquired it is stored in the short
term memory. In order for it not to be forgotten it has to be consolidated
into the long term memory. The capacity of the short term memory to
store is very limited but the capacity of the long term memory bank is
much larger.

Different informational aspects of long term memory traces seem to be processed and
coded, then stored in conjunction with other memories of the same type. It takes longer
time to retrieve information from the LTM because the
stores are larger. Remembering is the process by which we retrieve specific
information from memory stores. Forgetting is the inability to retrieve
stored information. Information in the STM is permanently forgotten while in the
LTM forgetting is only temporary. Memory traces are present in
multiple regions of the brain. The neurons involved are widely distributed
throughout the cortical and sub cortical regions of the brain. The regions
of the brain implicated in memory include the temporal lobes, the
prefrontal cortex, other regions of the cerebral cortex, the limbic system
and the cerebellum.

The temporal lobes and limbic system are essential for transferring new
memories into long term storage. The hippocampus plays a vital role in
integration of various related stimuli in the STM. It is also crucial for consolidation
into LTM. However the hippocampus stores only temporarily new LTM and then
transfers them to other cortical structures for more permanent storage. Accessing and
manipulating these long-term stores appears to be carried out by the prefrontal region of
cerebral cortex. The cerebellum seems to play a role in procedural memories involving

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motor skills gained through repetitive training. The hippocampus and surrounding
regions are responsible for declarative memory. Amnesia (loss of memory) has been
found to result from damage to the temporal lobe of the cerebral cortex, hippocampus,
head of the caudate nucleus, or dorsomedial aspects of the thalamus.

2.6 Midbrain
The mesencephalon or midbrain is located between the diencephalon and
the pons. It connects the pons and cerebellum with the cerebrum
(forebrain). On the ventral surface of the midbrain is a pair of cerebral
peduncles, made up of pyramidal tracts (fibres to the motor nuclei of the
spinal nerves within the spiral cord) corticobulbar (motor fibres to the
cranial-nerve motor nuclei) and corticopontine fibres to the pons. The third
cranial nerve (occulomotor) emerges from the fossa between the
peduncles on its ventral side. Passing through the midbrain is the cerebral aqueduct. The
dorsal portion of the midbrain situated above the aqueduct is the roof which has 4 little
elevations - the colliculi.

The colliculi are known collectively as the corpora quadrigemina. The superior pair of
colliculi are reflex centres that coordinate the movements of the eyeballs and head,
regulate the focusing mechanism in the eyes and adjust the size of the pupils in
response to visual stimuli. Cranial nerve IV (trochlear) emerges from the
roof of the midbrain. The inferior colliculi just posterior are relay nuclei of
the auditory pathways going to the thalamus and eventually to the auditory
cortex.

The midbrain also contains the red nucleus, an area of grey matter deep in the midbrain.
It maintains connections with the cerebrum and cerebellum and is involved in motor
coordination. Another nucleus is also present, a heavily black pigmented nucleus called
substantial nigra. It is integrated into neural circuits with the basal ganglia and therefore

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is involved also with motor-coordination (somatic motor activities).The substantial nigra
has a role in Parkinson's disease.

2.7 The Hindbrain


The hindbrain (rhombencephalon) is composed of the metencephalon and the
myeloencephalon which we will discuss separately.

2.7.1-Metencephalon
The region is composed of the pons and the cerebellum. The pons can be
seen as a bulge on the underside of the brain, between the mid brain and
the medulla oblongata. The pons is composed mainly of fibres that connect the
hindbrain to the midbrain as a relay station. Surface fibres in the pons connect to the
cerebellum, and sensory tracts that pass from the medulla oblongata, through the pons
and onto the midbrain. Within the pons are several nuclei associated with specific cranial
nerves - Trigeminal (V) abducens (VI), facial (VII) and Vestibulocochear (VIII). Other
nuclei in the pons cooperate with nuclei in the medulla oblongata to control breathing.
Two respiratory control centres are in the pons - the apneustic and pneumotaxic
centres.

The cerebellum occupies the inferior and posterior aspect of the cranial
cavity and is the second largest structure of the brain. It contains outer
grey matter and inner white matter (like the cerebrum). Fibres from the
cerebellum pass through the red nucleus to the thalamus and then to the
motor areas of the cerebral cortex. Other fibre tracts connect the cerebellum with
the pons, medulla oblongata and spinal cord. The cerebellum receives inputs from
proprietors (receptors in joints, tendons and muscles) and working together with the
basal ganglia and motor areas of the cortex participate in the coordination of movements.

The cerebellum consists of three functionally distinctive parts with different functions:

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1. The Vestibulocerebellum is important for the maintenance of balance and control of
eye movement.
2. The Spinocerebellum regulates muscle tone and coordinates skilled, voluntary
movements. It receives signals from the cortex concerning specific message to muscles
and also receives inputs from peripheral receptors concerning body movement and
position. The spinocerebellum essentially acts as "middle management"
comparing the "intentions" or "orders" of the higher centres with
the "performance" of the muscles and correcting deviations.
3. The Cerebrocerebellum plays a role in planning and initiation of voluntary activity by
providing input to the cortical motor area. It is also the region involved in procedural
memories. Damage or disease of the cerebellum produces the following range of
symptoms which reflect the loss of the aforementioned function:- poor
balance, nystagmus, reduced muscle tone but no paralysis, inability to
perform rapid movement smoothly and inability to stop and start skeletal
muscle action quickly. All these are due to a lack of coordination due to
errors in speed, force and direction of movement, a condition called ataxia.
The condition is also characterised by intention tremor which occurs only
when intentional movements are made. The person may reach for an
object and miss it by overshooting or placing the hand too far to the left or
right of the object, and then attempt to correct it by repeating the to and
from movement. This back and forth movement can result in oscillations of the limb.

2.7.2 Myeloencephalon
This is made up of only the medulla oblongata. The medulla measures
about 3cm long and is continuous with the pons superiorly and the spinal
cord inferiorly. The medulla, with the pons and midbrain make up the
brain stem. All the descending and ascending tracts that provide communication
between the spinal cord and the brain must pass through the medulla.
The vertical surface of the medulla contains bilateral elevated ridges called

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the pyramids. These pyramids are composed of the fibres of motors
tracts from the motor cerebral cortex to the spinal cord. These fibres
(pyramids) cross to the contralateral (opposite) side at the lower part of the
medulla to the opposite side of the spinal cord, forming an "X". This crossing over of
motor nerve fibres is called decussation. The result is that
the left side of the brain receives sensory information from the right side
of the body and vice versa. Similarly, the right side of the brain controls
motor activity in the left side of the body and vice versa.

Many important nuclei are contained in the medulla. Some of them are
involved in motor control while some of them form nerve roots for many
of the cranial nerves. Cranial nerves IX, X, XI and XII all have rootlets in
parts of the medulla. The vagus nuclei located one on each lateral side of
the medulla give rise to the highly important vagus (X) nerve. Other nuclei
are there which relay sensory information to the thalamus and them to the
cerebral cortex. The medulla also contains groupings of neurons that make up the vital
centres. These include the cardiac (cardioinhibitory centre) for the parasympathetic
inhibition of the heart, the vasomotor centre, for control of the autonomic innervations of
blood vessels, and the respiratory centre which acts together with centres in the pons to
control breathing.

2.8 Reticular Formation


Running throughout the brain stem and into the thalamus and hypothalamus is a
widespread and complex network of neurons and fibres called the reticular formation. It
is organised into (1) ascending (sensory) pathways from ascending spinal cord tracts
and the cerebellum, (2) descending (motor) pathways from the cortex and
hypothalamus; and (3) cranial nerves. Ascending fibres from the reticular formation carry
signals upwards to arouse and activate the cerebral cortex. These fibres compose
the reticular activating system (RAS) which controls the overall degree of cortical

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alertness and the ability to direct attention. Because of its many interconnections, the
RAS is activated in a non-specific fashion by any modality of sensory information.
Not surprisingly, general anaesthetics may produce consciousness by depressing the
RAS. Also our ability to fall asleep may be due to the action of specific
neurotransmitters that inhibit activity of the RAS.
ITQ 2: Enumerates the lobes of the cerebrum
ITA 2: four (4) major ones and probably an additional minor fifth lobe. The lobes are the frontal,
parietal, temporal and occipital lobes, which are visible from the surface and deep and fifth insula
lobe

3.0 Conclusion/Summary
We have been studying how the brain is a very important component of the central
nervous system which is responsible for controlling and coordinating almost all
activities of the human organisms. I have also tried to show you how these functions are
carried out by the use of an enormous number of neurons, associating with one another in
various regions and sub-regions of the brain. I am sure you have also learnt that:
I. The brain is structurally organised through the embryonic neural tube, from where the
structures developed upwards moving from the hindbrain up to the forebrain. The
direction of sophistication and consciousness also follows the direction of development.
By the 5th - 6th week of intrauterine life the brain had differentiated into 5 regions which
are still used to indicate the general regions of the brain as follows: (1)
telencephalon (cerebral cortex) and (2) diencephalon both of which make
up the forebrain. (3) mesencephalon which is the midbrain(4) metencephalon (which
consists of the pons and cerebellum) and (5) myeloencephalon
which contains the medulla oblongata. These last two make up the
hindbrain. The adult brain contains up to 100 billion neurons.

2. The cerebrum consists of two hemispheres connected by a large fibre tract called the
corpus callosum. The outer part of the cerebral cortex is grey matter and underneath is
white matter; however there are still nuclei of grey matter buried within the white matter
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of the cerebrum called basal nuclei. The two cerebral hemispheres have a degree of
specialisation of functions termed cerebral lateralisation, but there is cooperation in the
functions of the two hemispheres aided by the corpus callosum. Particular regions of the
cortex (left hemisphere) appear to be more important in language ability.

3. The limbic system and the hypothalamus are brain regions implicated as centres
for various emotions. Memory traces involve numerous brain regions but the
hippocampus of the medial temporal lobes in particular appears to control
consolidation of short term memory into long term memory.

4. The diencephalon is the region of the forebrain that includes the epithalamus,
thalamus and hypothalamus. These structures control many functions in the body as
well as serve as important relay centres for sensory information.

The structures of the midbrain and hindbrain make up the brainstem which
performs many functions. The brain stem serves as a connecting link between the rest
of the brain and the spinal cord. Most of their functions are concerned with incoming
and outgoing fibres traversing between the periphery and the higher brain centres with
incoming fibres relaying sensory information to the brain and outgoing ones carrying
command signals from the brain for efferent output.

4.0 Self-Assessment Questions


1. Describe the structural organization of the brain
2. Explain the anatomical divisions of the cerebrum and their functions
3.Describe the function of the midbrain
4. Explain the function of the Reticular formation
5. Identify the various nuclei within the white matter of the brain.

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5.0 References/Further Readings
Fox, S.I. (1996). Human Physiology Boston: Wm. C. Brown Publishers.
Carola, R. Harley J.P., and Noback C.R. (1990). Human Anatomy and
Physiology. New York: McGraw Hill.
Sherwood L. Human Physiology: From Cells to Systems. Minneapolis:
West Publishing Co.

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STUDY SESSION 2
The Spinal Cord, Cranial Nerves and Spinal Nerves
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 - Review of the Functional Anatomy of the Spinal Cord
2.2-Functional Role of the Spinal Cord: White Matter
2.3-Functional Role of the Spinal Cord: The Grey Matter
2.4-Functional Role of the Spinal Cord: The Spinal Reflex
2.5- Cranial Nerves
2.6-Spinal Nerves
3.0 Conclusion/Summary
4.0 Self-Assessment Questions
6.0 References/Further Readings

Introduction:
After the brain which we studied in the previous session, the spinal cord is the second
part of the central nervous system that extends through the vertebral canal and is
connected to spinal nerves. The spinal cord is strategically located between the brain
and the peripheral nervous system; and this location enables it to fulfil its primary
functions. One of the primary functions of the spinal cord is that it serves as a link for
transmission of information between the brain and the remainder of the body. In this
unit, we will examine the physiology of the spinal cord. We will also look at cranial and
spinal nerves.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. review briefly the anatomy of the spinal cord
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2. discuss the functional roles of the spinal cord in terms of the following:
- White matter
- Gray matter
- The spinal reflex
3. Describe the 12 cranial nerves and functions
4. Describe the spinal nerve and their functions

2.0 Main Content


2.1 Review of the Functional Anatomy of the Spinal Cord

The spinal cord is the part of the central nervous system that extends from the level of the
foremen magnum of the skull downward for about 45cm to the level of the first
lumber vertebra (L1) in adults. The upper end is continuous with the lowermost part of
the brain (medullar). Its lower end tapers off as the cone-shaped conus terminalis,
located in the vicinity of the first lumber vertebra. The spinal cord is encased in the bony
vertebral column which serves to protect the cord. The central opening or foramen in the
vertebral column has a diameter about the size of your index finger. The cylindrical cord
inside the column is about as thick as a pencil, or ones’ little finger. Paired
spinal nerves emerge from the spinal cord through the spaces formed
between the bony, wing like arches of adjacent vertebrae. The nerves are
named according to the region of the vertebral column from which they
emerge.

There are 31 pairs of spinal nerves and roots out of which there are 8 cervical (C)
nerve pairs, 12 thoracic (T), 5 lumber (L), 5 sacral (S) and one coccygeal (Co). Each pair
of spinal nerves passes through a pair of intervertebral foramina between two successive
vertebrae and distributed to a specific pair of segments of the body. The roots of all nerves
passing caudally below the conus resemble flowing coarse strands of hair. Therefore the
bundle of fibres from the lumber and sacral roots is called canda equina (horse tail)

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because of its appearance. The spinal cord and nerve roots are also protected by the
spinal meninges (membranes) and cerebrospinal fluid. 3 layers of meninges cover the
brain. The outer layer is called the dura mater and is a tough fibrous membrane.
The middle layer is the arachnoid mater delicate and transparent. The innermost layer
is the tender pia mater, thin highly vascularised and tightly attached to the spinal cord.

Cerebrospinal fluid (CSF) is a clear watery ultra-filtrate solution formed


from blood. In the capillaries of the brain the fluid passes through small
openings from the 4th ventricle into the subarachnoid spaces around the
brain and spinal cord and returns to the blood. CSF provides a cushion
which protects the delicate tissues of the cord. A cross-section for the
spinal cord shows that the anatomy is generally the same throughout its
length. Unlike in the brain where grey matter forms a cover over the white
matter, the grey matter of the spinal cord is located centrally, surrounded
by white matter. The inner grey matter is arranged to form a butterfly (or H) shaped
region with two dorsal horns and two ventral horns. The white matter is composed of
ascending and descending fibre tracts which are arranged into six columns called funiculi.
The spinal cord is divided into more or less symmetrical halves by a deep groove called
the anterior median septum and the posterior median septum.
The two primary functions of the spinal cord are:
i. To serve as a link for transmission of information between our brain and the
remainder of our body and
ii. To integrate reflex activity between afferent input and efferent output without
involving the brain.

ITQ 1: What are the various membranes covering the spinal cord?
ITA 1:
-Dura mater
- Arachnoid mater
- Pia matter

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2.2 Functional Roles of the Spinal Cord (The White Matter)
The white matter of the spinal cord surrounds the inner butterfly shaped
region of grey matter. The white matter is composed mainly of myelinated
nerve fibres and myelin has whitish colour. The white matter is divided into three pairs
of columns (funiculi) of myelinated fibres running the entire length of the cord. The
funiculi consist of the anterior (ventral) column, the posterior (dorsal) column and the
lateral column. The bundles of fibres within each funiculus are organised as tracts. A
tract is a bundle of nerve fibres (axons of long interneuron) with a similar function. Each
tract begins or ends in a particular area of the brain (depending on its type).
Each transmits a specific type of information. Some types are
ascending tracts (cord to brain) while others are descending tracts (brain to
cord). The tracts are named to indicate their location whether they are ascending
or descending, and their origin and termination. Ascending tracts usually
begin with spine and end with the brain region where the fibres first
synapse. E.g. the lateral spinothalamic tract originates in the grey matter of
the spinal cord synapses at the thalamus before being relayed to the
cerebral cortex and is located laterally in the spinal cord. Descending tracts
begin with a prefix showing the brain region where they originate and end
with the suffix-spinal. E.g. corticospinal tracts begin in the cerebral cortex
and descend the spinal cord.

2.2.1 Ascending Tracts


Ascending tracts are made up of sensory fibres that carry impulses up the spinal cord to
the brain. They convey sensory information from cutaneous receptors, propioceptors
(muscles and joint senses) and visceral receptors. Most of the sensory information that
originates in the right side of the body crosses over to eventually reach the region on
the left side of the brain for analysis. Similarly, the information arising in the left side
of the body is analysed by the right side of the brain. The decussation occurs in the
medulla oblongata for some sensory modalities or it can occur in the spinal cord for

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other modalities of sensation. Some sensory pathways (tracts) have sequences that are
made up of three neurons. In some path ways (tracts) the neurons in the sequence are
called first, second and third order neurons. A first order neuron extends from the
sensory receptor to the CNS; a second-order neuron extends from the spinal cord
or brain stem to a nucleus in the thalamus and a third-order neuron goes from the
thalamus to the sensory area of cerebral cortex.

Table 3.2.1: The principal ascending tracts of the spinal cord

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2.2.2 Descending Tracts
Descending tracts of motor fibres transmit impulses from the brain down the spinal
cord to the efferent neurons. The motor (descending) tracts and associated neural
circuits are grouped into two as pyramidal (corticospinal) and extra pyramidal
tracts.

The pyramidal tracts descend directly to the spinal cord. The cell bodies
that have their fibres in these pyramidal tracts are located primarily in the
precentral gyrus (motor cortex). Other areas of the cerebral cortex, however, also
contribute to these tracts. About 80% of the corticospinal fibres decussate in the pyramids
of the medulla oblongata (hence the name) and descend as the lateral corticospinal
tracts. The remaining fibres that did not decussate form the
anterior possibly ventral corticospinal tracts. These ones decussate in the
spinal cord. Because of the crossing over of the pyramidal tract, the right
cerebral hemisphere controls the musculature on the left side of the body
and the left hemisphere control the right musculature. The corticospinal
tracts are primarily concerned with the control of fine movements that
require dexterity.
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The remaining descending tracts are extra pyramidal motor tracts. They
originate in the midbrain and brain stem regions. Many centres, complex
circuits within the brain and several descending pathways are involved.
The system includes all the pathways that influence and regulate the motor
control of the lower motor neurons except those of the pyramidal tracts.
The regions of the cerebral cortex, basal nuclei and cerebellum that
participate in this motor control have numerous synaptic interconnections
and can influence movement only indirectly by means of stimulation or
inhibition of the particular nuclei that give rise to the extra pyramidal tracts. If the
pyramidal tract is cut in an experimental animal, stimulation
of the cortex, cerebellum and basal ganglia can still produce movements.

The major tracts in the extra pyramidal system are the reticulospinal tracts. They originate in
the reticular formation of the brain stem which receives either stimulatory or inhibitory
input from the cerebrum and cerebellum. This is because there are no descending tracts
from the cerebellum. The cerebellum can only influence motor activity indirectly by
affecting the vestibular nuclei, basal nuclei etc.

Summary of Descending Motor Tracts to Spinal Interneuron and Motor Neurons


Tract Category Origin Crossed or Uncrossed
1. Lateral corticospinal Pyramidal Cerebral Crossed cortex
2. Anterior corticospinal Pyramidal Cerebral Uncrossed cortex
3. Rubrospinal Extra pyramidal Red nucleus Crossed (midbrain)
4. Tectospinal Extra pyramidal Superior Crossed culliculus (midbrain)
5. Vestibulospinal Extra pyramidal Vestibular Uncrossed nuclei (in medulla oblongata)
6. Reticulospinal Extra pyramidal Brain stem Crossed reticular formation (medulla and
pons)

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2.3 Functional Role of the Spinal Cord: The Gray Matter
The centrally located grey matter is also functionally organised. The central canal
filled with CSF lies in the centre of the grey matter. The butterfly shaped grey matter
has two horns, the dorsal (posterior) and the ventral (anterior) horns. The two
posterior horns function in afferent input while the anterior horns function in efferent
somatic output. The pair that forms the cross bar of the H shape is known as the grey
commissure. It functions in cross reflexes. The dorsal horncontains cell bodies of
interneurons where afferent neurons with sensory input synapse. The ventral horn
contains cell bodies of neurons that supply skeletal muscles. Spinal nerves connect
with each side of the spinal cord by a dorsal root and a ventral root.

Afferent fibres carrying in coming signals enter the spinal cord through the dorsal root;
efferent fibres carrying outgoing signals leave through the ventral root. Groups of cell
bodies whose axons make up the dorsal root lie outside the cord, and are called dorsal
root ganglia (A ganglion is a collection of cell bodies outside the CNS; in the CNS they
are called centre or nuclei). The cell bodies of the efferent neurons (called anterior
horncells) originate in the grey matter of the cord and their axons make up the ventral root.
The dorsal and ventral roots at each level join up to form a spinal nerve that emerges
from the vertebral afferent and efferent fibres traversing between a particular region of the
body and the spinal cord.

2.4 Functions of the Spinal Cord (The Spinal Reflex)


A reflex is any response that occurs automatically without our conscious
effort. There are two types of reflexes, namely, simple or basic reflexes, which are
built in unlearned responses, such as closing the eyes when an object
moves towards them; and acquired or conditioned reflexes, which are a
result of practice and learning. An example of this is a pianist striking a
particular key on seeing a given note on the music staff. The musician
does this automatically, but only after considerable conscious training. The neural pathway
involved in accomplishing reflex activity is known as
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a reflex arc. The reflex arc typically includes five basic components as
follows:
1) Receptor
2) Afferent pathway
3) Integrating centre
4) Efferent pathway
5) Effector

The receptor detects a stimulus as a physical or chemical change in the


environment of the receptor. In response to the stimulus, the receptor
produces an action potential that is relayed by the afferent pathway to the
integrating centre which is usually in the CNS. (The spinal cord and
brain are responsible for integration of basic reflexes. The integrating
centre processes all information available to it from this receptor and all
other inputs; then makes a decision about the appropriate response. This
instruction is transmitted through the efferent pathway to the effector (a
muscle or a gland which carries out the required response). Unlike conscious
behaviour, a reflex response is predictable because the pathway
between the receptor and effector is always the same.

A basic spinal reflex is one integrated by the spinal cord i.e. all
components necessary for linking afferent input to efferent response are
present within the spinal cord. An example is the withdrawal reflex. For example, when you
touch a hot object, a reflex is initiated to withdraw your hand
from the hot object. Receptors are present in our skin for hot, cold,
warmth, light touch etc. The information is sent as action potentials by the
afferent system to the CNS. Once the afferent neuron enters the spinal
cord, (integrating centre) it diverges to synapse in different ways. One of
the ways is to stimulate an excitatory interneuron that will stimulate the
efferent motor neuron which supplies the biceps to flex the elbow and

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therefore withdraw the hand.

It may stimulate inhibitory interneuron which would antagonise the desired response e.g.
to inhibit the triceps from extending the elbow. The third is to stimulate other interneuron
to carry the message up the spinal cord to our brain where the impulse is appropriately
interpreted for what it is, its location and where it can be stored as memory and we can
start thinking about what to do about
it. However, a spinal reflex can be modified by the brain to override the input from the
receptors; actually preventing the biceps from contracting in spite of the painful stimulus.

Fig. 3.2.1: Spinal reflex

2.5 Cranial Nerves


There are twelve pairs of cranial nerves which are the peripheral nerves of
the brain. Two of these pairs arise from the forebrain and ten pairs arise from the midbrain
and hindbrain. The cranial nerves are designated by Roman numerals and by names. The
Roman numerals refer to the order in which the nerves are positioned from the front of the
brain to the back. The names indicate structures innervated by these nerves (e.g. facial) or
the principal function of the nerves (e.g. oculomotor). Cranial nerves are classified as
sensory, motor and mixed. The sensory fibres are afferent fibres associated with the special

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senses. The cell bodies of these sensory neurons are not located in the brain but are found
in ganglia near the sensory organ.
The motor fibres of the cranial nerves emerge from the brainstem. They arise from bundles
of neurons called motor nuclei, which are stimulated by nerve impulses from many outside
sources including the cortex of the cerebrum and sense organs. Axons of motor cranial
nerves have two roles. They (1) stimulate voluntary muscles or (2) Synapse with ganglia of
the autonomic nervous system which then relay nerve impulses to cardiac muscle, smooth
muscle and glands. The rest of the cranial nerves are mixed nerves. This term indicates
that the nerves contain both sensory and motor fibres. The cranial nerves are concerned with
the specialised senses of smell, taste, vision, hearing and balance and the general senses
and other inputs.

They are also involved with the specialised motor activities of eye
movement, chewing, swallowing, breathing, speaking and facial
expression. The vagus nerve is an exception projecting fibres to organs in the
abdomen and thorax.

Table 3.2.2: Summary of Cranial Nerves


Nerve Type Origin/Distribution Function
i. Olfactory Sensory Nasal Mucous membrane to Olfaction
olfactory bulb of cerebrum
Ii Optic Sensory Retina of the eye terminates in Vision, Afferent limb
lateral geniculate body of of reflex of focusing
thalamus and superior culliculus and constricting pupil.
of midbrain
iii. Oculomotor Motor Midbrain to all extrinsic muscles Movement of eyeball,
of eyeball except superior oblique elevation of upper
and laterial rectus; also autonomic eyelid constriction of
fibres to lens ciliary muscles of pupil focusing of lens.
lens and constrictor muscles
iv. Trochlear Motor Caudal Midbrain superior oblique Eye movements (down
muscles of eye. and out)
v. Trigemical (Opthalmic Sensory Pons to general area of forehead, Conveys general sense
division) eyes. from cornea of yeball

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upper nasal cavity front
of scalp forhead, upper
eyelid conjutiva,
lacrimal gland
vi. Maxillary division Sensory Pons to general area of maxillary Conveys general sense
region from cheek upper lips
upper teeth, mucosa of
nasa cavity, palate,
parts of pharynx
vii. Mandibular Mixed Pons sensory branch to area of Sensory conveys
mandibular region motor: general senses from
inervvates muscles of mastication tongue (not taste)
lower teeth skin of
lower jaw, motor
chewing.
viii. Abducens Motor Caudal pon innervate lateral Abduction of eye
rectus muscle of the eye (Lateral movement)
movement.
ix. Facial Mixed Pons sensory innervates taste Sensory taste
buds Motor: of tongue motor:
salivation,
innervates muscles of
lacrimation, facial expression,
movement of
autonomic fibres to facial muscle
salivaryglands, lacrimal glands
x. Vestibulocochlear Sensory Auditory: medulla, Hearing r Vestibular:
pons to cochlear of equilibrium Medulla, pons to
inner ear. Semicircular ducts
utricle and saccule of
inner ear
xi. Glosso-Pharyngeal Mixed Medulla Sensory: Motor: innervates
Secretion of conveys taste from stylo-pharyngeus,
saliva, posterior third of muscle autonomic
swallowing tongue, general fibres stimulate parotid
senses from upper pharynx. gland
xii. Vagus Mixed Medulla to voluntary Medulla to voluntary
Swallowing, muscles of soft Swallowing,
palate, monitoring cardiac muscles of soft palate,
muscle, oxygen and monitoring
smooth muscle in carbon dioxide cardiac muscle, oxygen
respiratory, level in blood; and smooth muscle
in carbon dioxide
respiratory, level in

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blood;
cardiovascular, sense
blood digestive
systems pressure,
other visceral activities
of affected systems.
xiii. Accessory (Spinal Motor Medular, Cervical Voice Medular, Cervical
Accessory) Production spinal cord to Voice Production
muscles(larynx); muscles of spinal cord to
larynx stemocleidosense; muscles(larynx);
movement mastoid, trapezius of muscles of larynx
ahead, shoulders stemocleidosense;
movement mastoid,
trapezius of ahead,
shoulders
xii. Hypoglossal Motor Medulla to tongue muscles Movements of tongue
during, speech,
swallowing muscle
sense.

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2.6 Spinal Nerves
We each have thirty one (31) pairs of spinal nerves. These nerves are grouped
according to the region of the vertebral column as follows:
8 cervical nerves
12 thoracic
5 lumber
5 sacral
1 coccygeal

Each spinal nerve is a mixed nerve composed of sensory and motor fibres.
These fibres are packaged together in the nerve, but separate near the
attachment of the nerve to the spinal cord. This produces two roots to each
nerve. The dorsal root is composed of sensory fibres and the ventral root
of motor fibres. The dorsal root contains an enlargement called the dorsal
root ganglion where the cell bodies of sensory neurons are located.
However, the cell bodies of the efferent (motor) fibres that innervate
skeletal (somatic) muscles are not in a ganglion but instead in the grey
matter of the spinal cord. The cell bodies for some autonomic motor neurons are
located in ganglia outside the spinal cord.

A short distance after the dorsal and ventral roots join together to form the spinal
nerve proper, the nerve divides into several branches called rami (singular
ramus) as follows the dorsal, ventral, meningeal ramui and the rami
communicants. Branches of the dorsal ramus innervate the skin of the back and
back of the head and the tissues and deep muscles of the back. Branches of the
ventral ramus innervate the skin, tissues and muscles of the neck, chest,
abdominal wall, both pairs of limbs and the pelvic area. The meningeal ramus
innervates the vertebrae spinal meninges and spinal blood vessels. The rami
communicants are composed of sensory (general visceral afferent) and motor
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fibres associated with the autonomic nervous system. The ventral rami of the
spinal nerves with the exception of T2 to T12 are arranged to form several
complex networks of nerves called plexuses. In a plexus the nerve fibres of
different spinal nerves are sorted and recombined. Plexuses include:
i. The cervical
ii. The branchial
iii. The lumber
iv. The sacral (sometimes the lumber and sacral plexus) and
v. The coccygeal.

The ventral rami of T2 to T12 do not form plexus; each ramus innervates a segment
of the thoracic and abdominal walls.

Figure 3.2.3 Diagram of Cranial nerves

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Table 3.2.4 Summary Spinal Nerve Plexus

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ITQ 2: Enumerate 31 pairs of spinal nerve
ITA 2:
1. 8 cervical nerves
2. 12 thoracic
3. 5 lumber
4. 5 sacral
5. 1 coccygeal nerve

3.0 Conclusion/Summary
In this session, we have been able to establish the following:-
1. The spinal cord is the neuronal link between the brain and spinal cord which
serves as the integrative centre for spinal reflexes. The spinal cord is the second part
of the central nervous system that connects the brain to the peripheral neurons
system.
2. A cross-section of the spinal cord show a butterfly-shaped centrally placed grey
matter and an outer layer of white matter.
3. The central grey matter of the spinal cord arranged in the form of H-shape has
two dorsal horns and two posterior horns which form the dorsal and ventral roots
of the spinal nerves.
4. The white-matter of the spinal cord is composed of ascending and descending
fibre tracts arranged into six columns.
5. The simple reflex is an unconscious motor response to a sensory stimulus
and this reflects a very important function of the spinal cord.
6. Twelve pairs of cranial nerves arise from the forebrain, midbrain and hindbrain
and they are concerned with specialised sensory and motor activities.
7. There are 31 pairs of spinal nerves. Each pair contains both sensory and motor
fibres. The dorsal root of a spinal nerve contains sensory fibres. The ventral root of
a spinal nerve contains motor nerves.

4.0 Self-Assessment Questions


1. Enumerate the 12 pairs of cranial nerves.

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2. With the aid of a well diagram describe the spinal reflex.
3. Identify land marks of the spinal cord.

5.0 References/Further Readings


Fox, S.I. (1996). Human Physiology Dubuque: Wm C. Brown Publishers.
Carola, R. Harley, J.P. of Noback, C.R. (1990) Human Anatomy &
Physiology. New York: McGraw Hill.
Sherwood L. (1993). Human Physiology from Cells to Systems.
Minneapolis:

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STUDY SESSION 3
Neurons and Neuronal Transmission
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
2.0 Main Content
2.1 Description of Neurons
2.2 Structural and Functional classification of Nervous
2.3 Transmission of Impulses
2.4 Action Potentials
2.5 Synapses
2.6 Transmission across a Synapse
2.7 Neurotransmitters
2.8 Neurotransmitters and their Receptors
2.9 Drugs and Synaptic Transmission
3.0 Conclusion/Summary
4.0 Self-Assessment Questions and Answers
5.0 Additional Activities (Videos, Animations & out of Class activities)
6.0 References/Further Readings

Introduction:
In this session, we will be looking at neurons. Our nervous system contains
about a trillion neurons and a lot more gill cells. Neurons are the basic building
blocks of the nervous system which is one of the two major regulating systems of
the body. The regulatory function of the nervous system is exerted over our
body’s muscular and glandular activities most of which are directed toward
maintaining homeostasis in our body.

Neurons are specialised for rapid integration and transmission of nerve

165
impulses. They are able to initiate, process, code and conduct changes in their
membrane potential as a means of transmitting a message rapidly throughout
their length. They can also transmit this information through intricate nerve
pathways from neuron to neuron, or neuron to muscles and glands through
chemical means.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. Describe the neuron and explain structural and functional classification of
neurons.
2. Explain the process of impulse transmission.
3. Describe an action potential showing the ionic basis of excitation and
conduction.
4. Describe the functional anatomy of a synapse.
5. Explain neuronal transmission across a synapse.
6. Explain the mechanism of action of neurotransmitters, their
inactivation, removal and recycling.
7. Explain the action of different neurotransmitters in the nervous system.

2.0 Main Content


2.1 Description of Neurons
Neurons are highly specialised cells that transmit impulses within animals to cause
a change in a target cell such as a muscle effector cell or glandular cell. Neurons,
together with neuroglial (glial) cells make up the nervous system. The neuron is
the integral element of our five senses and other physical, regulatory and mental
faculties including memory and consciousness.
Neurons have three main purposes:
1. To gather and send information from the senses such as touch,
smell, sight etc.

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2. To send appropriate signals to effector cells such as muscles,
glands etc.
3. To process all information gathered and provide a memory
and cognitive ability thus allowing us to take
action on information received.
The structure of neurons is essentially the same in all animals, although
the human nervous system is much more specialised and complicated than
that of lower animals. Neurons are divided into different regions each
having a different function. A typical neuron has the following parts:

1. A cell body (soma) which contains the cell nucleus and the
cytoplasmic organelles which are responsible for maintaining the cell
2. Several short processes called dendrites which are radial extensions of cell
membrane of the body which extend to other neurons and increase the surface area
available for connecting with other axons from other neurons
3. A long process extending from the soma, called the axon. The axon
can sometimes stretch over a very long distance and is responsible for
transmitting signals from the neuron to other cells downstream in the chain.
The axon divides into terminal branches each ending in a button-like
structure called synaptic knobs or terminal buttons.
4. Specialised cell junctions called synapses between an axon and other
cells which allow for direct communication from one cell to another.

Neuroglial or neurilemma cells are associated with axons of neurons. In


the central nervous system they are called oligodendrocytes and in the
peripheral nervous system, they are the Schwann cells. They associate
with the axons by wrapping themselves around portions of the axon. This
forms a segmented sheath around the axon called myelin sheath. Myelin is
a lipid (fatty) substance which is pale in colour. It is this colour that leads
to the term white matter or grey matter within the nervous system. The
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white matter contains myelinated axons predominantly whereas the grey
matter contains mainly neuronal cell bodies which are non-myelinated.
The function of the myelin is to increase the electrical capacitance of
neurons and to insulate against any leakage of nerve impulse. They also
assist in nourishing the axon. The higher the capacitance, and the better the
insulation, the faster the nerve impulse will travel along the axon.

The gap of about 1mm between segments of myelin sheath (junctions


between adjacent neuroglial cells) is known as nodes of Ranvier. These serve
important functions in neural transmission of impulses in a very specialised
way.
Neurons display a very high level of metabolic activity for some reasons:
i. They have massive surface areas compared to other cells in the body.
ii. They need to generate nerve impulses.
iii. Their high level of activity reflects on the appearance of the cell— the
nucleus is large, with prominent nucleolus representing a high degree
of cellular activity.
iv. There is abundant endoplasmic reticulum for protein synthesis.
v. There is a well developed Golgi apparatus to provide secretions
especially neurotransmitters.
vi. There are also large numbers of mitochondria to produce the large
amount of energy required by the neuron.

The function of the nervous system as a whole depends very much upon the
complexity of the network of connections between the various neurons rather
than the specific features of any single neuron.

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2.2. Structural and Functional Classification
Neurons exist in many shapes and sizes and their structure affects their
functions. The structural classification of neurons depends on the number
of processes extending from the soma. Multipolar neurons that have many
dendrites and usually have one long axon carrying an impulse away from the cell
body. They also tend to have large cell bodies. Majority of the neurons in the
spinal cord are multipolar and they serve principally as motor neurons.

Bipolar neurons have only two main processes similar in length a single dendrite
and an axon. Bipolar neurons are generally small simple cells that provide local
connections within the central nervous system. They can also be found in some
sense organs i.e. retina and olfactory cells. Unipolar neurons possess one major
process which subdivides into two branches; one running to the CNS (axon)
and the other to a part of the body (dendritic in function). They are usually
sensory neurons.

Figure 3.3.4: The diagram of a Neuron

Functionally neurons are classified as sensory neurons, motor neurons or


interneurons. Sensory neurons transform physical stimuli (sensations) such as

169
smell, light or sound into action potentials which are then transmitted to the
central nervous system. They always have their dendrites in a sensory receptor
in the body and the axons end within the central nervous system (either the
spinal cord or the brain depending on the part of the body in which the
sensory receptor is located). Since they send impulses to the central nervous
system, sensory neurons are also called afferent neurons. Motor neurons
transmit nerve impulses away from the brain and spinal cord to muscles or
glands and are also called efferent neurons. Interneurons, also called
association neurons provide local connections within the central nervous
system. They are mostly found in the central nervous system and are
responsible for receiving, relaying integrating and sending nerve impulses
within the CNS. All neurons whatever their structure or function carry nerve
impulses in one direction. That is from dendrite to cell body and from cell
body to the axon. Dendrites always carry impulses toward the cell body
and axons always carry impulses away from the cell body.

2.3 Transmission of Nerve Impulses


The transmission of nerve impulses which is the unique function of nerve cells
depends on two properties of nerve cells:

Excitability and Conductivity


Excitability refers to the fact that nerve cells are able to respond to a
stimulus, and in fact have a threshold for excitation. The stimulus may be
internal or external; electrical, chemical or mechanical. Muscle fibres are
also able to demonstrate excitability and this makes them able to contract
when stimulated.

Conductivity refers to the property that neurons alone possess which


makes them able to transfer their excitability along their length and then on to

170
other neurons or even muscle tissue.

These two properties together allow neurons to deliver appropriate messages


to appropriate parts of the body as and when required.

All cells in our body possess a membrane potential which is related to the
non-uniform distribution of and differential permeability to Na+, K+ and
large intracellular anions (protein ions). Nerve cells and muscle cells have
specialised use for this membrane potential. They are able to undergo
transient, rapid changes in their membrane potential i.e. they can alter their
transmembrane potential reversibly because of the differences in the ionic
concentration inside and outside the cell and the selective permeability to
Sodium (Na+) and potassium (K+). We say that nerve and muscle cells are
excitable tissues because of their ability to produce electrical signals when
excited.
In a resting neuron, the concentration of potassium (K+) inside the cells is
up to 30 times greater than it is outside whereas the concentration of
sodium is up to 14 times less inside than outside. Even in this state of non-
conduction of nerve impulses the neuron continuously maintain a balance
of sodium-potassium pump across the membrane i.e. Na+ are actively
transported out and K+ are actively transported into the cell. The inside of
the cell at rest is negatively charged because K+ are though positively
charged and are abundant within the cell, there are a large number of
negatively charged (protein) ions within the cell which cannot diffuse out
through the cell membrane. The K+ is not enough to balance out the
protein anions within the cell and this leads to an overall negative charge
within the cell. Outside the cell, the larger quantity of Na+ with no protein
anion leads to an overall positive charge in the extracellular fluid. This state
of potential difference in polarity across the membrane is known as the resting
membrane potential. It is usually about-70mV.
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When a nerve cell is stimulated one of two types of physiochemical
disturbance is produced: local, non-propagated potentials called
electrotonic potentials or a propagated disturbance called action potential or
nerve impulses. All excitable tissues exhibit these electrical responses and
these are the main language of the nervous system. When an excitable tissue is
sufficiently stimulated, the polarity of the membrane is reversed momentarily
with the inside becoming more positive compared to the outside. It is this
event (shift) that we refer to as the action potential.

2.4 The Action Potential


The stimulation of an excitable tissue represents the delivery of energy to the cell
membrane in some way. For example, the energy from sound waves exciting the
neurons in the inner ear; heat energy exciting neurons in the skin etc. Action
potentials are brief reversals of membrane potential brought about by rapid
changes in membrane permeability occasioned by physiochemical
stimulation. Within nerve cells the action potential has an amplitude of
approximately +110mv (millivolts) and lasts for about 1 millisecond (MS).
Action potentials follow the all-or-none law by which we mean that they are not
graded responses but are either full-sized or absent depending on the strength of
the stimulus. i.e. if the stimulus is strong enough the action potential is fired at
its full strength of + 110mv. If the stimulus is not strong enough, there is no action
potential at all. The action potential is also able to spread throughout the
membrane in non-decemental fashion. Four terms are normally used to describe
the processes that occur during action potentials:

Polarisation: This refers to the fact that the membrane has potential
(difference) i.e. there is a separation of opposite charges with negative charges
inside and positive charges outside. This is the state of the membrane at rest.

172
Depolarisation: This refers to the stage when the membrane potential is
reduced from resting potential; it has decreased or moved toward zero. There
is a mix up of opposing charges; only few charges are separated. It is shown
as an upward deflection on a recording device.

Hyperpolarisation: This refers to when potential difference has returned and


increased or become even more negative than -70mV. A downward
deflection is shown on the measuring device.

Depolarisation: The membrane potential returns to its normal resting


value. To initiate an action potential there is a triggering event that causes the
membrane to depolarise. Depolarisation normally proceeds slowly at first
until it reaches a critical level known as threshold potential about 55mv.
After this initial 15 mV of depolarisation, an explosive increase in
depolarisation occurs. There is a sharp upward deflection past the zero
level to up to +30 to +35mv. This is called the overshoot because the
deflection overshoots the zero potential. This is followed again by a rapid
decrease towards OmV and a fall rapidly toward the resting level, a
process called repolarisation. Often the repolarisation to resting potential
is driven back too far causing a brief period of hyperpolarisation (below -70mv).
This period lasts about 40ms, before the membrane finally returns to its resting
potential (repolarisation).

The action potential fails to occur if the stimulus is sub-threshold in


magnitude whereas if the stimulus is at threshold or above threshold
intensity it occurs with constant amplitude and form regardless of the
strength of the stimulus. (The all or none law). Although sub-threshold
stimuli do not produce an action potential, they do have an effect on the
membrane potential. When there is a triggering event of sub-threshold

173
level, there is localised depolarising potential charge. This change can rise
very sharply but also drops off rapidly as it moves away from the initial
source of stimulation. These graded potentials or electrotonic potentials
function as signals only for short distances and die out as the distance from
the initial source of stimulation increases.
2.4.1-Basis of Excitation and Conduction
The nerve cell membrane as has been discussed is polarised at rest with
positive charges lined up along the outside of the membrane and negative
changes along the inside. During the action potential this polarity is abolished
and for a brief period actually reversed. The cell membranes of nerves like those
of other cells contain different types of ion channels or gates. Some are
passive (continually open) and some are voltage-gated (i.e. open or close in
response to changes in membrane potentials) while some are chemical
messenger-gated (ligand-gated). These open or close in response to the
binding of a specific chemical messenger for example neurotransmitter,
hormone, to a membrane receptor. The voltage-gated channels are the ones
mostly involved in action potentials.

At rest Na+ is actively transported out of the cell and K+ is actively


transported into the cell. However K+ diffuses out of the cell down its
concentration gradient and Na+ diffuses back in through their leak
channels (passive). However the permeability of the membrane to K+ is
much greater (because of the leak channels) therefore passive K+ efflux is
much greater than passive Na+ influx. The resting membrane is 50 to 75
times more permeable to K+. This in addition to the impermeability of the
membranes to most protein anions within the cell maintains the membrane
in a polarised state with the outside more positive than the inside (- 70mv).
When a membrane starts depolarising toward threshold as a result of a
triggering event, some of the voltage-gated Na+ channels open. Since the

174
concentration and electrical gradient of Na+ both favours its movement into
the cell, Na+ starts to move in, carrying its positive charges with it. This
depolarises the membrane further, thereby opening more Na+ gated
channels and allowing more Na+ to enter.

This continues in a positive feedback cycle. There is therefore an


explosive increase in Na+ permeability at threshold potential as the
membrane becomes 600 times more permeable to Na+ than to K+ Na+
rushes into the cell rapidly eliminating the internal negativity and
making the inside even more positive than the outside. This is
represented by the steep upward deflection on a measuring device, past the
zero potential level and up to an overshoot level of +30 to +35 mv. The potential
does not however become more positive or rise higher than this level. This is
because at this peak level of the action potential the Na+ channels slam shut
and enter a state called the inactivate state until the membrane potential has
even restored to its negative resting value a period of a few milliseconds.
In addition, the direction of the electrical gradient for Na+ reverses and this
limits the influx. The cell membrane becomes impermeable again. At the same
time, as the inactivation of the Na+ channels occurs, K+ permeability greatly
increases i.e. the voltage-gated K+ channels open and more K+ move out of the
cell. The opening of the K+ channels is slower but more prolonged than the
opening of Na+ channels and can actually be considered a response triggered by
the depolarisation caused by Na+ influx. The marked increase in K+ permeability
causes K+ to rush out of the cell down its concentration and electrical gradients
carrying positive charges back to the outside.

At the peak of an action potential the very positive inside of the cell also tend to
repel the positive K+ ions so that the electrical gradient for K+ favours outward
movement. The outward movement of K+ rapidly restores the internal
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negatively returning the potential to resting. This completes the repolarisation
process and is shown by the downward deflection past the zero potential
towards the initial resting potential of -70mv on the measuring device. Because
the K+ channels do not close very quickly, more K+ can actually leave the cell
than is necessary. This slightly excessive K+ efflux makes the interior of
the cell more negative than resting potential; and this explains the after
hyperpolarisation period.

At the end of an action potential, the membrane potential has been restored
to its resting condition but the ion distribution had been slightly altered.
Sodium has entered the cell during the rising phase while a comparable
amount of K+ has left during the falling phase. It is now left for the Na+ -
K+ pump to restore these ions to their original locations in the long run.
Depolarisation of one part sends an electrical current to neighbouring un-
stimulated parts of the membrane, which stimulates adjacent portions of
the neuron membrane to also depolarise. This event repeats itself along the
membrane of the cell, thereby conveying the nerve impulse along the
neuron. This continuous conduction is the way impulses are transmitted
down unmyelinated nerve fibres. In myelinated nerves, the myelin sheath
forms an insulating layer around the axon therefore depolarisation can
only occur at the nodes of Ranvier where there are short sections of non-
myelination. Impulses are conducted by sequential jumping from one node
to another along the nerve.

This form of impulse conduction is usually quicker than continuous


conduction. It is known as saltatory conduction.

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2.5 Synapses
Impulses are transmitted from one nerve cell to another cell at synapses. A
synapse is a junction where the axon of one neuron (the pre-synaptic
neuron) meets the dendrite, cell body or even axon of another neuron or in
some cases, a muscle or gland cell (the post synaptic neuron or cell).

2.5.1-Functional Anatomy of Synapses


A neuron may terminate at one of three structures; another neuron, a
muscle, or a gland. Specifically the use of the word at this point will be
limited to the junction between two neurons. The commonest synapse is
formed at the junction between an axon terminal and the dendrites or cell
body of a second neuron. Less frequently axon to axon and dendrite to
dendrite connections occur. Usually most neuronal cell bodies and their
dendrites receive thousands of synaptic input (axon terminals) from many other
neurons

Figure 3.3.5: Diagram of a synapse (inset)

177
The axon terminal of a pre-synaptic neuron is usually slightly enlarged to
form button-like structures called the synaptic knobs. The synaptic knobs
contain numerous synaptic vesicles which store a specific chemical
messenger, called a neurotransmitter. These neurotransmitters have been
synthesised by the presynaptic neuron. The membrane of the synaptic
knobs (of the presynaptic neuron) does not come into direct physical
contact with the membrane of the post-synaptic cell, rather a small gap of
about 20 nanometers separate the two. This space is called the synaptic
cleft. This space makes it difficult for electrical signals (action potentials)
to pass directly between the two cells; the presynaptic and the post
synaptic neurons-thereby necessitating the action of a mediator. In most
synapses transmission is by chemical means requiring the chemical
mediators neurotransmitters. At some synapses however transmission is
electrical while in a few synapses it is conjoint i.e. both chemical and
electrical being possible.

In any case, transmission across a synapse is not


a simple jumping of action potentials from the presynaptic to the
postsynaptic cell. Synapses operate in one direction only i.e. the presynaptic
neuron brings in the signal to the synapse and stimulates the postsynaptic neuron
which then carries signals away from the synapse.

When an action potential in a presynaptic neuron reaches the synaptic knob, it


triggers the opening of the voltage-gated calcium ion (Ca++) channels in the
synaptic knobs. This allows Ca++ which is more in the extracellular fluid (outside
the cell) to enter inside the cell. The presence of Ca++ inside the cell causes the
vesicles inside the knob to move towards the cell membrane at the synaptic cleft.
Vesicles fuse with the membrane and release their neurotransmitters by
exocytosis into the synaptic cleft. The neurotransmitters diffuse across the space
178
and bind to receptors in the membrane of the post-synaptic cell. The amount of
neurotransmitter released is proportionate to Ca++ influx. The binding of
neurotransmitters to receptors in the postsynaptic cell membrane opens ion
channels in the post-synaptic cell membrane. This is an example of a chemical
messenger-gated channel. With the opening of the channels there is a change in
the permeability of the post synaptic membrane, resulting in a change in the
membrane potential. If the excitation is strong enough it results in the generation
of a nerve impulse.

The neurotransmitter molecules left in the synaptic cleft are usually broken
down by enzymes, reabsorbed by pre-synaptic cell where they are
resynthesised (using energy from ATP generated by the nearby
mitochondria) and packaged once again into vesicles.
2.6.1-Synaptic Excitation and Inhibition
We have two types of synapses depending on the permeability changes
induced in the post synaptic neuron by the combination of transmitter
substances with receptor sites: excitatory synapses and inhibitory
synapses. At an excitatory synapse, the response to the receptor
neurotransmitter combination is an opening of the Na+ and K+ channels in
the post synaptic membrane, increasing the permeability to both of these ions. Na+
moves into the cell in large numbers, reducing the potential difference in
the cell by making the inside of the cell less negative than at rest. This
produces a small depolarisation of the post-synaptic neuron. The effect of just one
synapse is usually not enough to stimulate the post-synaptic membrane to a
firing level (threshold). Before a post-synaptic membrane
can fire an action potential, it must be stimulated by several synapses at
once. Each synapse produces a small depolarisation and each depolarisation
contributes to bringing the membrane nearer to threshold and increase the
likelihood of firing an action potential.

179
This post synaptic potential change occurring at an excitatory synapse is called an
excitatory post-synaptic potential (EPSP). The small rises contributed by
depolarisations at each synapse add together, raising the EPSP to a level high
enough to trigger a nerve impulse (fire an action potential). We refer to it as
spatial summation when stimulation occurring at the same time at several
synapses add together to reach threshold level. However, if the same synapse
supplies impulses to the post-synaptic neuron in quick succession before the
previous ones have died out, the stimulations add together in what is called
temporal summation. Certain synapses however have opposite effects. The
effect of the neurotransmitter receptor combination instead of opening the
Na+ channels rather opens the K+ or Cl- channels.

The result is that the membrane becomes hyperpolarised and the inside of
the post synaptic membrane becomes more negative. What happens is that if it
is the K+ channels that open, more positive charges leave the cell via K+
efflux, leaving more negative charges behind inside the cell or in the case of
increased Cl- permeability negative charges enter the cell in the form of Cl-
ions because Cl- concentration is higher outside the cell. The slight
hyperpolarisation moves the membrane potential even farther away from
threshold making it more difficult for excitation to threshold level to
occur. The membrane is said to be inhibited under these circumstances,
and the small hyperpolarisation is called inhibitory post-synaptic potential
(IPSP). Both spatial and temporal summation of inhibitory potentials can
also occur. The transmission of an electrical signal across a synapse, for a
presynaptic neuron takes time, an interval of at least 0.5ms (0.5 - Ims). This is
called synaptic delay, and corresponds to the time it takes for the
neurotransmitter to be released and to act on the post-synaptic membrane.

Therefore it is faster to transmit electrical signals through a pathway with


fewer synapses than a complex pathway with multiple synapses. The more
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complex (polysynaptic) the pathway, the more synaptic delays and
therefore the longer the time required to respond to a particular electrical
event. The generation of an action potential (nerve impulse) in the post-synaptic
neuron is therefore the result of a constant interplay of excitatory and
inhibitory activity of thousands of presynaptic neurons on the postsynaptic
neuron. This produces a fluctuating membrane potential i.e the algebraic
sum of the hyperpolarising and depolarising activity. The cell body of the
neuron performs an integrating function. When the net effect of excitatory
and inhibitory activities at synapses produces 10 - 15mv of depolarisation, which is
sufficient for an action potential to fire, an action potential results
i.e an impulse is transmitted.

2.7 Neurotransmitters
We know or suspect that many different chemicals act as neurotransmitters
in the nervous system. Neurotransmitter substances vary from synapse to synapse
and the same transmitter is always released at a particular synapse. One particular
neurotransmitter will always induce EPSP while another will always induce
IPSP. Yet another neurotransmitter may even induce an EPSP in one synapse and
an IPSP in another synapse. The response of a given transmitter B receptor
combination at a given synapse is always constant. A given synapse is either
always excitatory or always inhibitory.

In our nervous system, some neurotransmitters are simple chemical


ions such as calcium, others are more complex chemicals such as
Dopamine, Serotonin (SHT), gamma-amino-butyric acid (GABA),
Acetyloline, etc. Identified neurotransmitters can be divided into broad
categories based on their chemical structure. Some are amines e.g.
dopamine, Norepinephrine, epinephrine etc, others are amino acids, e.g.
glutamate, aspartate, glycine etc, others are purines e.g. adenosine, and many

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are polypeptides e.g. somatostatin, Endothelins, Endorphins, motilin,
Glucagon, Gastrin, angiotensin II etc. Some of these substances in addition to
acting locally as neurotransmitters in the synaptic cleft can also function as
hormones at other sites distant from where they are produced or act as paracrine
regulators. It is also known that many neurons contain more than one transmitter
i.e. they contain co transmitters. Often the amines exist with one or more
polypeptides.
2.7.1-Neurotransmitter Inactivation, Removal and Recycling
Neurotransmitters are quickly inactivated from the synaptic cleft once it
has produced the appropriate response in the post-synaptic neuron so that
the post-synaptic neuron can get ready for other presynaptic inputs. As
long as neurotransmitters remain bound to their receptors, EPSP or IPSP
which they produce continue, thus, it is necessary that they are removed
and their responses terminated. We can divide this removal in the
following ways:
They may be inactivated by specific enzymes within the membrane of the
post-synaptic neuron or they may be actively taken up back into the axon
terminal by transport mechanisms in the presynaptic membrane
(reuptake). Once the neurotransmitter is inside the synaptic knob,
(following reuptake) it can either be (1) Stored and released another time
(recycling) or (2) Destroyed by enzymes in the synaptic knob.

2.8 Neurotransmitters and Their Receptors


Receptors are protein substances on the surface of the cell or in some instances
in the cytoplasm or nucleus which act as binding sites for chemical
messengers (hormones, neurotransmitters and other ligands). There are three
facts that must be noted about receptor.
1. Every ligand (chemical messenger e.g. neurotransmitter) has many sub types of
receptors. For example, nor epinephrine binds to α1, α2, ß1 and ß 2 receptors. There

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are also different kinds of α1 and α2 receptors. This makes the possible effects of
particular lagans more specific and more selective.
2. Receptors can exist on the presynaptic as well as post-synaptic neurons. The
presynaptic receptors usually act to inhibit further secretion of the ligand (by
feedback control). For example noradrenaline binding to α2 presynaptic receptors
can inhibit norepinephrine secretion.
3. The subtypes of receptors tend to group in large families as far as structure and
function is concerned. Thus, some families of receptors in combination with their
lagans function by changing the lagan-gated channels thereby altering membrane
permeability and ionic fluxes across the postsynaptic membrane. Another mode of
synaptic transmission used by the transmitter-receptor complex
involves the activation of second messengers within the postsynaptic neuron
such as cyclic AMP which can then perform the function of opening the ion
channels or other functions as may be necessary.

2.8.1 Acetylcholine as a Neurotransmitter and its Receptors


Acetylcholine is an amine neurotransmitter that exists commonly and in
high concentrations in terminal buttons of cholinergic neurons. Neurons
which release acetylcholine are known as cholinergic neurons.
Acetylcholine receptors are divided into two main types due to their
pharmacologic properties:- muscarinic and nicotinic receptors depending on
the action of acetylcholine on the different parts where they function.
Muscarine has stimulatory action on smooth muscles and glands, thus the
muscarinic actions of acetylcholine are stimulatory on smooth muscles and
glands. The receptors here are called muscarinic receptors. They are blocked by
the drug atropine.

In autonomic ganglia, large amounts of acetylcholine block transmission of


impulses from pre- to post-ganglionic neurons. These are nicotine-like actions.

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Thus these actions of acetylcholine are nicotinic actions, and the receptors for
such actions are called nicotinic receptors. Acetylcholine nicotinic receptors have 5
sub-units:- two alpha, one beta, one gamma and one delta subunits. Acetylcholine
binds on alpha subunits and when it does, it opens ionic channels for Na+ and
other cat ions, resulting in the influx of Na+ and a depolarising potential.
Muscarinic receptors also have four types identified and they seem to act
through a second messenger system. Acetylcholine is removed from the
synaptic cleft through the catalytic activity of acetylcholinesterase which
hydrolyses acetylcholine to choline and acetate.

2.8.2 Catecholamine and their receptors


Norepinephrine is the neurotransmitter found at most sympathetic
ganglionic endings. Together with its methyl derivative epinephrine they
are secreted by the adrenal medulla, however epinephrine is not a mediator
at preganglionic sympathetic nerve endings. The neurons secreting
norepinephrine are called adrenergic neurons. Sometimes, the term adrenergic
is used to refer to both of them. The third catecholamine in the body is
dopamine and dopamine secreting neurons are called dopaminergic
neurons. The catecholamines are formed by hydroxylation and decarboxylation of
the amino acids phenylalanine and tyrosine. Epinephrine and norepinephrine both
act on alpha and ß receptors, with norepinephrine having greater affinity for X
adrenergic receptors and epinephrine for ß adrenergic receptors. Both α and ß
receptors work through the action of cyclic AMP as second messenger.

Dopamine which is a step in the formation of norepinephrine and


epinephrine (catecholamines can be secreted as a neurotransmitter in
certain parts of the brain and autonomic ganglia. There are the D1 and D2
receptors at sites where dopamine is released. Their action is by activating
dopamine-sensitive adenylate cyclase (second messenger system). Catecholamines

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are recaptured by active reuptake mechanism and inactivated by monoamine
oxidase (MAO) and catechol-O-methyltransferase (COMT). Other classical
neurotransmitters of the amine type include serotonin, and histamine.

2.8.3-Amino Acids and polypeptides as neurotransmitters


Other transmitters of the amino acid and polypeptide group differ from the
classical transmitters in that they are larger molecules containing sometimes
from two to forty amino acids. They also bring about slower
and more prolonged responses. Some of these substances released at
synapses function as true neurotransmitters but some as neuromodulators.
Neuromodulators are chemical messengers that bind to neuronal receptors
at non-synaptic sites and by so doing bring about long-term changes that
depresses or enhances effective synaptic transmission, often by activating
second messenger systems.

For example a neuromodulator may influence the level of an enzyme critical


in the synthesis of a neurotransmitter by a presynaptic neuron or alter the
sensitivity of a post synaptic neuron to a neurotransmitter.

2.9 Drugs and Synaptic Transmissions


Many drugs are able to interfere with neurotransmitter processes and this has
resulted in their use as legitimate drugs for treatment of mental health problems or
as illegal or recreational drugs.
The vast majority of drugs that influence the nervous system perform their function
by altering synaptic mechanisms. Hence they can be used to block an
undesirable effect or to enhance a desirable one. Specifically some of those
drugs may act in the following ways:
i. Altering the synthesis, axonal transport, storage or release of a
neurotransmitter;

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ii. Modifying neurotransmitter interaction with the post synaptic
iii. receptor,
iv. Influencing neurotransmitter reuptake or destruction; or
v. Replacing a deficient neurotransmitter with a substitute transmitter.

For example, the illegal drug cocaine blocks the reuptake of the
neurotransmitter dopamine at presynaptic terminals by competitively
binding with the dopamine reuptake transporter which picks up released
dopamine from the synaptic cleft and shuttles it back to the axon terminal
for reuptake. When cocaine occupies the dopamine transporter, dopamine
remains longer in the synaptic cleft, and continues to interact with its
receptor sites in the post synaptic neuron. This results in prolonged
activation of the neural pathways that use dopamine as neurotransmitter,
such as those pathways that play a role in emotional responses like
feelings of pleasure.

Drugs that alter neurotransmitter functions are called psychoactive drugs.


They can be divided into six major pharmacological classes based on their
desired behavioural or psychological effect: alcohol, sedative hypnotics,
opiate analgesics, stimulant euphoriants, hallucinogens, and psychotropic agents.
Hallucinogens are pscychedelic drugs such as LSD (Lysergic acid
diethylamide), mescaline etc. they are serotonin agonists that produce their effects
by activating and binding to 5HT2 receptors. They are usually taken
illegally to alter perception and thinking patterns. They have little known
medical use. Psychotropic drugs like the phenothiazine tranquilisers are effective in
the relief of symptoms of schizophrenia. Their antipsychotic activity parallels their
ability to block D2 dopamine receptors.

2.10 Disease and Synaptic Transmission


Synaptic transmission is also vulnerable to a number of disease processes
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including defects at presynaptic and post-synaptic sites. Parkinson’s disease for
example is attributable to a deficiency of dopamine in a particular region
of the brain involved in controlling complex movements called the
substantial nigra. The cells have their axons ending in the basal nuclei of the
brain. A gradual destruction of the dopamine secreting cells in the substantial nigra
and the resultant loss of basal nuclei function are responsible for Parkinson’s
disease. The basal nuclei is another region of the brain involved in the
coordination of slow, sustained movements, inhibition of muscle tone and
suppression of useless patterns of movement.

As dopamine activity slowly diminishes, symptoms begin with involuntary


tremors at rest, such as involuntary rhythmic shaking of hands and head.
Symptoms of increasing stiffness and rigidity ensue as disease worsens. Treatment
of Parkinson’s disease is an example of a deficient neurotransmitter being replaced
with a substitute transmitter. Patients with this disease are given Levidopa (L-
dopa) a closely related precursor of dopamine. The drug can be taken up by the
dopamine deficient synaptic knobs, thereby substituting for the lacking, naturally
occurring dopamine. It alleviates the symptoms associated with dopamine deficit.

Strychnine and tetanus toxin act at different synaptic sites to block


inhibitory impulses while leaving excitatory inputs untouched. Strychnine
competes with the inhibitory transmitter glycine at the post synaptic
receptor sites. It takes up receptor sites without affecting the cells
potential, making the receptors not available for binding with glycine
when it is released. In such nerve pathways that use glycine as
neurotransmitter, post-synaptic inhibition is abolished. Unchecked
excitatory pathways lead to convulsions and muscle spasticity. Tetanus
toxins also prevent the release of another inhibitory transmitter, Gamma
aminobutyric acid (GABA) from presynaptic inputs terminating on motorneurons

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supplying skeletal muscles.

Unchecked excitatory inputs to these neurons result in uncontrolled muscle


spasms. The outcomes of the two are similar, but strychnine blocks specific
postsynaptic inhibitory receptors whereas tetanus toxin prevents the presynaptic
release of a specific inhibitory neurotransmitter. Other drugs and diseases that
affect synaptic transmission are too numerous to mention but the examples
show that any site along the synaptic pathway can be interfered with
pharmacologically or pathologically.

ITQ 1: Define the concept of a synapse


ITA 1: A synapse is a junction where the axon of one neuron (the pre-synaptic neurone)
meets the dendrite, the cell body or even axon of another neuron or in some cases, a muscle
or gland cell (the post synaptic neuron or cell)

3.0 Conclusion/Summary
In this session, I believe we have sufficiently dealt with neurons and how nerve
cells are specialised to receive, process, encode and rapidly transmit information
from one part of the body to another. The information is transmitted over intricate
nerve pathways by propagating action potentials along the nerve cells length as
well as by chemical transmission of the signal from neuron to neuron and later
from neuron to muscle or gland through neurotransmitter - receptor
interactions at synapses. The specialisation of nerve cells depends on their
ability to rapidly alter their membrane potential and thus produce electrical
signals in response to appropriate triggering event. In addition, we have also learnt
that:
i. Neurons are highly specialised cells which together with neuralgia
cells make up the nervous system; and they function mainly to gather
and process information from parts of the body to the central nervous

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system and to bring back appropriate information from the nervous
system to relevant parts of the body (usually muscles and glands).
ii. Structurally, neurons are classified as multipolar bipolar and unipolar
reflecting the arrangement of its process (axon and dendrites).
iii. Functionally, they can be classified as afferent or efferent reflecting the
direction of conduction of impulses in relation to the central nervous
system. They are also functionally classified into sensory motor, or
interneurons.
iv. The unique function of impulse transmission is made possible in the
neurons by their two properties of excitability and conductivity. Nerve
cells are excitable tissues because they can alter their trans membrane
potential (reversibly) due to selective permeability of their membranes
to Na+ and K + . This excitation can also be propagated down the entire
length of the nerve cell. This is conduction.

v. An action potential is an electrical event in a nerve cell characterised by


brief reversals of membrane potentials brought about by rapid changes
in membrane permeability as a result of physiochemical stimulation.

vi. Action potentials (excitation and conduction) are made possible


because of the existence of ion channels through which electrical
charges can flow in and out of the cells. During an AP the permeability of
the membrane to Na+ and K+ is greatly altered permitting the flow of these
cat ions in and out of the cell. The concentration of these cations inside and
outside the cell determines the degree of polarisation in nerve cells.

vii. Nerve cells also have to transmit impulses across nerve to nerve
junctions called synapses. A neurotransmitter is required to carry the
electrical signal across the synapse.

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viii. Some synapses excite the post-synaptic neuron whereas others inhibit it.

ix. Neurotransmitters are quickly removed from the synaptic cleft;


inactivated and recycled in readiness for another use.
x. Receptors provide binding sites for neurotransmitters on the post-
synaptic membrane. he transmitter receptor complex functions by
altering membrane permeability. However some of them function
through intracellular second messenger system.
xi. The effectiveness of synaptic transmission can be modified by drugs and
diseases. Any site along the synaptic pathway is vulnerable to
interference.

4.0 Self-Assessment Questions


1………………………….. refers to the fact that the membrane has potential
(difference) i.e. there is a separation of opposite charges with negative charges
inside and positive charges outside. This is the state of the membrane at rest.
2………………………….. refers to the fact that membrane potential is
reduced from resting potential; it has decreased or moved toward zero. There
is a mix up of opposing charges; only few charges are separated. It is shown
as an upward deflection on a recording device.
3………………………….. refers to the potential difference has returned and
increased or become even more negative than -70mV. A downward deflection is
shown on the measuring device.
4. explain the process of impulse transmission
5. write a comprehensive note showing the ionic basis of excitation and
conduction of an action potential

190
7.0 References/Further Readings
Sherwood, Lauralee (1993). Human Physiology from Cells to Systems.
Minneapolis: West Publishing Co.
Hall J.E. (2016). Guyton and Hall textbook of medical physiology.
Philadelphia: Sainders Co.
http://www. jdaross.avc.net/intronerve 3.thitus Nov. 5:2007
http://normaly.sandhills.cc.nc.us/psy150/neuron.html Aug. 30 2005

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STUDY SESSION 4
The Autonomic Nervous System
Section and Subsection Headings:
Introduction
1.0 Learning Outcomes
4.1 Main Content
2.1 Structures and Pathways of the Autonomic Nervous System
2.1.1 Neural Control of Involuntary Effectors
2.1.2 Autonomic Neurons
2.2 - Visceral Effector Organs
2.2.1 - Divisions of the Autonomic Nervous System
2.2.1 - Sympathetic Division
2.2.2 - Parasympathetic Division
2.2.3 Functions of the Autonomic Nervous System
2.3 - Neurotransmitters of the Autonomic Nervous System
2.3.1 - Responses to Adrenergic Stimulation
2.3.2 - Responses to Cholinergic Stimulation
2.4 - Dual Innervations of Visceral Organs
2.5 -Control of Autonomic N/S by Higher Brain Centres
2.6-Somatic Nervous system
3.0 Conclusion/Summary
4.0 Self-Assessment Questions and Answers
5.0 Additional Activities (Videos, Animations & Out of Class activities)
7.0 References/Further Readings

Introduction:
Our focus now shifts to the autonomic nervous system which is considered to be
the involuntary branch of the peripheral efferent division. Skeletal muscles are
innervated by the somatic nervous system while cardiac muscle; smooth

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muscle most exocrine glands and some endocrine glands are innervated by
the autonomic nervous system. Two neurotransmitters: acetylcholine and
norepinephrine are released at the neuronal terminals and are responsible for
bringing about all the changes effected by the autonomic nervous system for
example bladder contraction and salivary secretion. The involuntary effects of
autonomic innervations contrast with the voluntary control of skeletal
muscles through the somatic neurons.

In this study session, we shall also be examining the structures and pathways of the
autonomic nervous system, the differences in the autonomic and somatic systems
as well as describe the structure and general functions of the sympathetic and
parasympathetic divisions of the autonomic system.

1.0 Study Session Learning Outcomes


After studying this study session, I expect you to be able to:
1. describe the structures and pathways of the autonomic system
2. explain the neural control of involuntary effectors
3. describe the sympathetic and parasympathetic divisions of
the autonomic nervous system
4. explain the functions of the autonomic nervous system list the
neurotransmitters in the autonomic nervous system and explain their
actions
5. explain the responses to adrenergic and cholinergic stimulation describe the
control of autonomic nervous system by higher brain centres
6. Briefly described the somatic nervous system comparing it with the
autonomic nervous system.

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2.0 Main Content
2.1 Structures and Pathways of the Autonomic System Compared to
Somatic System
As we have said in the Introduction, the autonomic nervous is the involuntary
branch of the peripheral nervous system. We can also refer to it as the visceral
efferent motor system because it is concerned with internal organs or viscera. The
autonomic nervous system is exclusively a motor system, involved with
influencing (innervating) the activity of cardiac muscle, smooth muscle and glands
of the body.

The autonomic nervous system consists of two divisions: the sympathetic


and the parasympathetic nervous system. Sympathetic nerve fibres originate
in the thoracic and lumber regions of the spinal cord. Each autonomic nerve
pathway extending from the CNS to an innervated organ consists of a two-neuron
chain. The cell body of the first neuron in the series is located in the CNS. Its
axon, the pre-ganglionic fibre, synapses with the cell body of the second neuron,
which the lies within a ganglion outside the CNS. The axon of the second neuron,
the post-ganglionic fibre, innervates the effector organ. The involuntary effects
of autonomic innervation contrast with the voluntary control of skeletal
muscles by way of somatic motor neurons.

ITQ 1: The autonomic nervous system consists of two divisions:


ITA 1: The sympathetic and the parasympathetic nervous system.

2.2 Neural Control of Involuntary Effectors


2.2.1 Autonomic Neurons
Neurons of the peripheral nervous system conduct impulses away from the central
nervous system. There are two major categories of motor neurons. Somatic motor
neurons have their cell bodies within the CNS and send axons to skeletal

194
muscles which are usually under voluntary control. Unlike somatic motor neurons,
which conduct impulses along a single axon from the spinal cord to the
neuromuscular junction, autonomic motor control involves two neurons in the grey
matter of the brain or spinal cord. The axon of this neuron does not directly
reach the effector organ but synapses with a second neuron within an
autonomic ganglion. The first neuron is called the preganglionic neuron while the
second in this pathway is called the post-ganglionic neuron and has an axon that
extends from the autonomic ganglion and synapses with the cells of an effector
organ. Preganglionic fibres originate in the midbrain, hindbrain and in the upper
thoracic to the 4th sacral levels of the spinal cord.

Autonomic ganglia are located in the head, neck and abdomen. Chains of
autonomic ganglia also parallel both sides of the spinal cord. The origin of the
preganglionic fibres and the location of the autonomic ganglia help to
differentiate the sympathetic and parasympathetic divisions of the autonomic
system.

2.2.2 Visceral Effector Organs


Since autonomic nervous system helps to regulate the activities of glands, smooth
muscle and cardiac muscle, autonomic control is an integral aspect of the
functioning of most body systems. Autonomic regulation therefore partly explains
the functioning of the systems/organs of the body like endocrine regulation,
functions of the heart and circulation etc. Unlike skeletal muscles which enter a
state of flaccid paralysis and atrophy when their motor nerves are cut, the
involuntary effectors are independent of their innervation. In fact, damage to an
autonomic nerve makes its target organs more sensitive than normal to
stimulating agents. Such compensatory mechanism may explain why the ability
of stomach mucosa to secrete acid may be restored after vagotomy.

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Smooth muscle and cardiac muscle have intrinsic muscle tone. In addition,
they can contract rhythmically even in the absence of nerve stimulation. This is
in response to electrical waves of depolarisation initiated by the muscles
themselves. Autonomic innervation simply increases or decreases this
intrinsic activity. Autonomic nerves also maintain a resting tone in the
sense that they maintain a baseline firing rate that can either increase or
decrease. The release of the neurotransmitter, acetylcholine from somatic
motor neuron always stimulates the effector organ (skeletal muscle). In contrast
some autonomic nerves release transmitters that inhibit the activity of their
effectors.

2.3. Divisions of the Autonomic Nervous System


The sympathetic and parasympathetic divisions of the autonomic system
have some structural features in common. Both consist of pre-ganglionic
neurons, which originate in the CNS, and post ganglionic fibres that
originate outside the CNS in ganglia. The specific origin of the
preganglionic fibres and their location are however different in the two
divisions.

2.3.1 Sympathetic Division (Thoracolumbar)


We can also call it the thoracolumbar division because its preganglionic fibres
leave the spinal cord from the first thoracic (TI) to the second lumber (L2)
levels. Most sympathetic nerve fibres however separate from the somatic motor
fibres and synapse with postganglionic neurons within a double row
of sympathetic ganglia or para vertical ganglia located on either side of the spinal
cord. The myelinated preganglionic sympathetic axons exit the spinal cord in
the ventral root of spinal nerves, but soon diverge from the spinal nerves within
white rami communicants. The axons within each ramus enter the
sympathetic chain of ganglia where they can travel to ganglia at different levels

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and synapse with post-ganglionic sympathetic neurons.

The axons of the post-ganglionic sympathetic neurons are unmyelinated and form
the grey rami communicates as they return spinal nerves to their effector
organ. Since sympathetic axons form a component of spinal nerves, they
are widely distributed to the skeletal muscles and skin of the body where
they innervate blood vessels and other involuntary effectors. Many preganglionic
fibres that exit the spinal cord in the upper thoracic level travel into the neck,
where they synapse in cervical sympathetic ganglia. From here post-ganglionic
fibres innervate the smooth muscles and glands of the head and neck. Many
preganglionic fibres that exit the spinal cord below the diaphragm
pass through the sympathetic chain of ganglia without synapsing.

Beyond the sympathetic chain of ganglia they form the splanchnic nerves. These
preganglionic fibres in the splanchnic nerves synapse in collateral ganglia.
These include the coeliac, superior mesenteric and inferior mesenteric
ganglia. Post ganglionic fibres that arise from the collateral ganglia
innervate organs of the digestive, urinary and reproductive systems. The adrenal
medulla, the inner portion of the adrenal glands is considered
a modified sympathetic ganglion, its cells having been derived from the
same embryonic tissue as ganglionic sympathetic neurons. It secretes the
hormones epinephrine (80%) and norepinephrine when stimulated by the
sympathetic system. Like a sympathetic ganglion, the preganglionic
sympathetic fibre enervates the adrenal medulla and causes it to secrete
epinephrine into the blood. The effect of epinephrine becomes comparable
to those of the neurotransmitter norepinephrine which is released at post
ganglionic sympathetic nerve endings. No other post ganglionic fibre is
needed. For this reason, and because the adrenal medulla is stimulated as
part of the mass activation of the sympathetic system, the two are grouped
together as a single sympathoadrenal system.
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Figure 3.4.1: Diagram of parasympathetic and sympathetic nerves

198
2.3.2 Parasympathetic (Cranio Sacral) Division
The preganglionic fibres of this division originate in the brain (specifically
midbrain, medulla, oblongata and pons) and in the second through 4th
sacral levels of the spinal column. It is therefore also called craniosacral division.
These fibres are long in comparison to sympathetic preganglionic fibres because they do
not end until they reach the terminal ganglia that lie next to or within the effector organs.
Most parasympathetic fibres do not travel within spinal nerves as do sympathetic
fibres. As a result, cutaneous effectors (blood vessels, sweat glands and erector pili
muscles) and blood vessels in skeletal muscles receive sympathetic but not
parasympathetic innervation.

Four of the twelve pairs of cranial nerves contain preganglionic


parasympathetic fibres. These are oculomotor (III) facial (VII)
glossopharyngeal (ix) and vagus (x) nerves. Preganglionic fibres from cell
bodies located in the midbrain are conveyed by the oculomotor (III)
cranial nerve to a synapse in the ciliary ganglion. The post ganglionic axon
terminals from there innervate the constrctor muscles in the iris, as well as
the ciliary muscles that change the shape of the lens to focus the eyes.
Preganglionic fibres originating in the lower pons leave by way of the
facial cranial (vii) nerve to either the pterygopalatine or submandibular
ganglia where they synapse. Post ganglionic fibres innervate the lacrimal
glands, which secrete tears, and the nasal, oral and pharyngeal cavities.
Preganglionic fibres of the glosso-pharyngeal nerve from nuclei in upper
medulla synapse in the otic ganglion which sends post ganglionic fibres to
innervate the parotid salivary glands.

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Preganglionic fibres that emerge from cell bodies located in the dorsal
vagal nucleus of the medulla are conveyed by the very long vagus (x)
nerve. They synapse in terminal ganglia located in many regions of the
body. The preganglionic fibres travel through the oesophageal opening in
the diaphragm into the abdominal cavity. In each region some of these pre-
ganglionic fibres branch from the main trunks of the vagus nerves and
synapse with postganglionic neurons located within the effector organs.
These very long preganglionic vagus fibres provide parasympathetic
innervation to the heart, lungs, oesophagus, stomach pancreas, liver, small
intestine and the upper half of the large intestine. Post-ganglionic
parasympathetic fibres arise from terminal ganglia within these organs and
synapse with effector cells (smooth muscles and glands). From the sacral levels of
the spinal cord arise preganglionic parasympathetic innervation to the lower half
of the large intestine, rectum and to the urinary and reproductive systems. These fibres
like those of the vagus synapse with terminal ganglia located within the effector organs.

2.4 Functions of the Autonomic Nervous System


The sympathetic and parasympathetic divisions of the autonomic nervous system
affect the visceral organs in different ways. Mass activation of the sympathetic
system prepares the body for intense physical activity in emergencies or stressful
situations, such as a physical threat from the outside environment. This response is
typically referred to as the flight or fight response because the sympathetic system
readies the body to fight against or flee from the threat. Let us take a moment to think
about the body resources needed in such circumstances. The heart beats more rapidly
and more forcefully; blood pressure is elevated because of generalised constriction of

200
the blood vessels; the respiratory airways open wide to permit maximal airflow,
glycogen (stored sugar) and fat stores are broken down to release extra
fuel in the blood; and blood vessels supplying skeletal muscles dilate.

The effects of parasympathetic stimulation are in many ways opposite to


the effects of sympathetic stimulation. They dominate in quiet, relaxed
situations. Under such non-threatening circumstances the body can be
concerned with its own general activities like digestion and emptying of
the urinary bladder. The parasympathetic system promotes these kinds of
bodily functions while slowing down those activities enhanced by the
sympathetic system. The parasympathetic system however is not normally activated as a
whole. Visceral organs respond differently to sympathetic and parasympathetic nerve
activity because the postganglionic fibres of these two divisions release different
neurotransmitters.

2.5 Neurotransmitters of the Autonomic Nervous System


Sympathetic and parasympathetic preganglionic fibres release the same
neurotransmitter, acetylcholine (ACH), but the postganglionic endings of
these two systems release different neurotransmitters. Parasympathetic
postganglionic fibres release acetylcholine. Accordingly, they, along with
all autonomic preganglionic fibres are called cholinergic fibres. In
contrast, most sympathetic post ganglionic fibres are called adrenergic
fibres because they release norepinephrine (noradrenaline). There are very
few exceptions where some sympathetic fibres release ACH. Examples
are in the sympathetic supply to blood vessels in skeletal muscles as well

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as to sweat glands.

2.5.1 Responses to Adrenergic Stimulation


It has been found that both excitatory and inhibitory effects can be
produced in different tissues by the same chemical. For example, adrenergic
stimulation from sympathetic nerves causes the heart, muscles of the iris and the
smooth muscles of many blood vessels to contract. However the same sympathetic
stimulation dilates the smooth muscles of the bronchioles and some blood vessels. The
possible explanation lies in the biochemistry of the tissue cells, especially differences in
the membrane receptor proteins. For example two major classes of these receptor
proteins have been designated alpha and beta adrenergic receptors. There are also
two subtypes of each class for example alpha 1 and alpha 2 and beta 1
and beta 2. From this, compounds which selectively bind to one or the
other of adrenergic receptor have been developed. As a result of their
binding capacity to adrenergic receptors, drugs have been developed
which either promote or inhibit adrenergic effect. It has also been possible
to determine which sub types are present in each organ.

A drug that binds to receptors for a neurotransmitter and promotes the


process stimulated by that neurotransmitter is called an agonist of that
neurotransmitter. A drug that blocks the action of a neurotransmitter is
said to be an antagonist. The use of drugs that selectively stimulate or
block a1, a2, ß1 and ß2 receptors has proved extremely useful in medical
application. For example, people with hypertension have been treated with
a beta-blocking drug known as propranolol. This drug blocks B1

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receptors to produce the desired effect of lowering the cardiac rate and
blood pressure etc.

2.5.2 Responses to Cholinergic Stimulation


Somatic motor neurons, all preganglionic autonomic neurons and most postganglionic
parasympathetic neurons are cholinergic releasing ACH as a neurotransmitter. The
cholinergic effects of somatic motor neurons are always excitatory. The cholinergic
effects of postganglionic parasympathetic fibres are also usually excitatory but there
are some notable exceptions. For example, the parasympathetic (cholinergic) effect on
the heart causes slowing of the heart rate instead of excitation. Just as adrenergic
receptors are divided into alpha and beta subtypes, cholinergic receptors are divided
into muscarinic and nicotinic receptor subtypes. The drug muscarine stimulates the
cholinergic receptors in the heart, digestive system, however does not stimulate
the muscarinic subtypes in autonomic ganglia or at the neuromuscular junction of
skeletal muscles. It is rather the drug nicotine that stimulates these cholinergic
receptors, therefore the receptors in these places must be nicotinic receptors.
The drug used for skeletal muscle relaxation blocks nicotinic receptors but has little
effect or muscarinic receptors.

2.6 Dual Innervation of Visceral Organs


Most visceral organs are innervated by both sympathetic and parasympathetic
nerve fibres. On a general note, the two systems exert
opposite effects in a particular organ. One system dominates at one time
while the other dominates other times depending on circumstances.
Usually both systems are partially active at each point in time until the

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particular circumstances that will cause the activity of one to dominate the
other ensue. These kinds of effect are called Antagonistic effects in conditions of dual
innervation. The best example of antagonism in the two systems is their
innervation of the pacemaker region of the heart. Here adrenergic
stimulation from sympathetic fibres increases the heart rate while
cholinergic stimulation from parasympathetic fibres decreases the heart
rate.

In a few cases the effects of sympathetic and parasympathetic nerves are


complementary or cooperative. Complementary effects occur when stimulation of
both divisions produce similar effects. An example is the sympathetic and
parasympathetic stimulation of the salivary glands. The effects are cooperative when
sympathetic and parasympathetic stimulation produce two different effects that work
together to promote a single action. An example is the parasympathetic effect on the
penis causing erection and the sympathetic effect producing ejaculation, they
cooperate to promote reproduction. A few organs in the body however do not have
dual innervation. These include the adrenal medulla, erector pili muscles, sweat
glands and most blood vessels which receive only sympathetic innervation.

2.7 Control of Autonomic Nervous System by Higher Brain Centres


Visceral functions are mostly regulated by autonomic reflexes, sensory input is
transmitted to the brain centres that integrate this information and respond by
modifying the activity of preganglionic autonomic neurons. The neural centres that
directly control the activity of autonomic nerves are influenced by higher brain centres
and by sensory input. The medulla oblongata in the brain stem is the area that most

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directly controls the activity of the autonomic system. Almost all autonomic responses
can be elicited by experimental stimulation of the medulla. The organ contains centres
for the control of cardiovascular, pulmonary, urinary, reproductive and digestive
systems.

The medulla oblongata itself is responsive to regulation by higher brain areas. One
of these is the hypothalamus which is the brain region that contains centres for the
control of body temperature, hunger, thirst, regulation of the pituitary gland and
together with the limbic system and cerebral cortex also controls various emotional
states. The limbic system which includes the cingulate gyrus of the cerebral
cortex, the hypothalamus, hippocampus and amygdaloidal nucleus is
involved in basic emotions like anger, fear, sex and hunger. The
involvement of the limbic system with the control of autonomic function is
responsible for the visceral responses characteristic of these emotional
states. Blushing, fainting, racing heart, cold sweats are examples of the
many visceral reactions that accompany emotions as a result of autonomic
activation.

2.8 Somatic Nervous System


The somatic nervous system is that part of the motor (efferent) division that supplies
skeletal muscles. The cell bodies of somatic motor neurons are located within the
ventral horn of the spinal cord. Unlike the two neuron chain of the autonomic system,
the axon of a somatic motor neuron is continuous from its origin in the spinal cord to
its termination on skeletal muscles. Motor neuron axon terminals release acetylcholine
which brings about excitation and contraction of the innervated muscles. The effect of

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motor neuron on skeletal muscles is only stimulation and never inhibition
or both. Inhibition of skeletal muscle activity can only be accomplished
within the CNS. Somatic motor neurons are influenced by many converging
presynaptic inputs both excitatory and inhibitory but the level of activity in a motor
neuron and its subsequent output to the skeletal muscle fibres depend on
the balance of EPSPs and IPSPs.

The motor neuron is considered to be the final common pathway by which any other
parts of the nervous system can influence skeletal muscle activity. The somatic
nervous system is considered to be under voluntary control but much of skeletal
muscle activity involving posture, balance and stereotypical movements are
subconsciously controlled.

ITQ 2: Describe the sympathetic nervous system


ITA 2: It is also called the thoracolumbar division because its preganglionic fibres leave the spinal
cord from the first thoracic (TI) to the second lumber (L2) levels.

3.0 Conclusion/Summary
Let us conclude this session by repeating that the CNS controls effector organs
(muscles and glands) by transmitting signals from the CNS to these organs through the
efferent division of the peripheral nervous system. The two parts of the efferent
system are the autonomic and somatic nervous system. Much of the efferent system
output is directed toward maintaining homeostasis in the body.

In summary,
1. The autonomic nervous system is the involuntary branch of the efferent motor
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system concerned with innervating and controlling the activities of cardiac muscle,
smooth muscle and glands.

2. The ANS is made of a two-neuron chain with preganglionic fibres originating in the
brain or spinal cord and postganglionic fibres originating in ganglia outside the CNS.

3. The ANS is divided into two systems: the sympathetic and Parasympathetic
systems. Preganglionic neurons in the sympathetic division originate in the spinal
cord, synapse with the postganglionic neurons located in a double chain of
paravertebral sympathetic ganglia outside the spinal cord. Others synapse at the
collateral ganglia. Some pregnaglionic fibres innervate the adrenal medulla which in
turn secretes hormones similar to the chemical transmitters from the sympathetic
postganglionic nerve endings.

4. Preganglionic parasympathetic fibres originate in the brain and sacral levels of the
spinal cord. They are long nerves synapsing with post ganglionic neurons within the
effector organs or very close to them.

5. The functions of the autonomic nervous system is evidenced in the functions of the
two major divisions. The sympathetic division exerts their effect through
adrenergic stimulation of effector organs. This chemical transmission activates the
body to flight or fight reaction necessary in emergencies.

6. The parasympathetic division mostly exert antagonistic effects to its counterpart


causing more relaxation and quiescence in the body and activating more routine bodily
functions like digestion.

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7. The body’s response to the two neurotransmitters of the ANS noradrenaline and
acetylcholine depends a lot on which receptor protein is present in each effect organ.
8. Higher brain centres like the medulla oblongata and the limbic system control the
autonomic nervous system.

9. The somatic nervous system is the part of the efferent peripheral nervous system that
innervates skeletal muscles.

5.0 Self-Assessment Questions


1. Describe the sympathetic and parasympathetic divisions of the
autonomic nervous system.
2. Explain the functions of the autonomic nervous system.
3. List the neurotransmitters in the autonomic nervous system and explain
their actions.

7.0 References/Further Readings


Carola et al (1990). Anatomy and Physiology. New York: McGraw B Hill.
Sherwood L. (1993). Human Physiology. West Publishing Company Minneapolis.

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XIII GLOSSARY

ablation Removal.

absorption Transfer of substances into the body or into the blood, usually across an epithelial
surface. E.g. absorption through the skin or from the gut, or re-absorption of substances from the
lumen of tubules in the kidney.

acclimatization, acclimation The process of adaptation to a new environment, for example with
altered ambient temperature, pressure or (with aquatic animals) salinity. Acclimation is sometimes
used specifically to mean adaptation to a single changed factor (as under laboratory conditions).

accommodation RELATED: adaptation ♦ A process of adjustment to new conditions. Used in


several senses: 1. Attenuation or absence of a response when a stimulus is applied slowly (e.g.
failure to detect a smell when it builds up slowly, or failure of a slow depolarisation to give an
action potential). NB this is distinct from adaptation, but is often seen in the same systems. 2.
Adjustment of focus of the eye.

acid CONVERSE: base ♦ An aqueous solution with a pH less than 7.0. A molecule or ion that can
dissociate to release a proton (H+). For example, H 2 CO 3 and H 2 PO 4 - are both acids.

acidosis CONVERSE: alkalosis ♦ Abnormally low pH of the blood. An acidotic condition may be
capable of being restored to normal acid-base balance by increasing respiratory ventilation to
reduce the CO 2 content of the body (respiratory acidosis), or it may require retention or injection of
HCO 3 - (metabolic acidosis).

acid-base balance Control of the factors influencing the pH of the blood. The main factors under
homeostatic control are the excretion of CO 2 , HCO 3 - and NH 4 +. Disturbances are lessened in the
short term by numerous pH buffer systems. Influences affecting blood pH are altered absorption,
metabolism and respiration, and disturbances affecting the control systems.

activation The process of initiating or increasing activity of some sort. In muscle physiology,
'activation' usually means the induction of action potentials in the cell membrane (= excitation)
rather than the initiation of tension (NB the link between the two is 'excitation-contraction
coupling').

active site The part of a cell or of a receptor that binds to a substrate.

active transport RELATED: carrier mediated transport ♦ Net movement of a substance across a
membrane from a lower to a higher concentration or (in the case of ions) against an
electrochemical gradient. This requires energy, and can occur by linkage of carrier mediated
transport to a process providing energy, such as the hydrolysis of ATP or the downhill movement
of another substance.

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adaptation Decline of a response while a stimulus is maintained constant after onset. Distinguish
carefully from both accommodation and habituation. Light and dark adaptation are the processes of
adjustment of the eye to different light levels (whether fast or slowly changing).

adequate stimulus The form of stimulation that normally elicits a specific reaction, such as an
action potential or a reflex response. ??

adrenaline, epinephrine (USA) A hormone released together with noradrenaline from the adrenal
medulla, especially in conditions of stress. Acts at both alpha and beta adrenergic receptors, and
mimics some of the effects of sympathetic nerve activation.

adult Sexually and/or emotionally mature.

aerobic CONVERSE: anaerobic ♦ Involving the use of oxygen. Aerobic conditions are ones in
which O 2 is present.

aetiology, etiology (USA) The cause of a disease.

afferent arteriole Arterioles carrying blood towards a structure organ, such as the kidney
glomeruli, in which there is arteriolar control of both afferent and efferent vessels.

affinity The strength of binding of two chemicals, or a chemical or ion to a receptor or enzyme.
Affinity of a receptor for an agonist may be expressed by the agonist concentration at which half of
the receptors are bound to agonist molecules (the dissociation constant for the complex: KD), or by
-log 10 of this quantity, or by its reciprocal.

agonist CONVERSE: antagonist ♦ Something that assists or mimics an action. For example,
agonist muscles pull parts of the skeleton in the same direction. Agonist drugs or chemicals bind to
the same receptors, producing the same effects.

alimentary Relating to food, or the gastro-intestinal tract.

all-or-none RELATED: threshold, positive feedback CONVERSE: graded ♦ A response that


appears only if the strength of a stimulus exceeds some threshold level, but whose amplitude and
characteristics do not depend on the stimulus, e.g. an action potential, or a sneeze. The sneeze is
more familiar: pepper either does or doesn't lead to a sneeze. A little pepper doesn't lead to a 'small'
sneeze, though it may lead to fewer sneezes. This isn't to say that sneezes are always identical: you
can stifle a sneeze and, for example, reduce the noise it makes. These alterations are nothing to do
with the stimulus that caused the sneeze, however. Exactly parallel things are true for action
potentials. All-or-none responses usually involve some form of 'positive feedback'.

altitude Height above sea level. This affects physiological processes largely through the reduced
atmospheric pressure, which falls exponentially by ca. 12% per 1000m rise (18% per 5000ft).
Oxygen percentage in inspired air remains constant (ca. 21%). Effects of altitude are often studied
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by simulation in a chamber with reduced pressure. Respiration is affected noticeably at 4000m
(13,000ft) and severely at 6000m (20,000ft). Reduced PO 2 leads to hyperventilation, which
partially maintains arterial PO 2 but lowers arterial PCO 2 .

alveolar air Air that is or has been in the alveoli. Samples of alveolar air can be taken at the end of
expiration after dead-space air has been expelled (end-tidal samples). Alveolar air is normally
approximately in equilibrium with arterial blood, and therefore has approximately the same partial
pressures of O 2 and CO 2 as arterial blood.

alveolus The terminal air sacs in the lungs, where gas exchange takes place with the blood.
Typically 150-300µm in diameter in man (ca. 5nl volume).

ambi- Both.

ambient Relating to the environment of an animal. E.g. ambient temperature, pressure. Note that
the conditions for a particular tissue may be different from the ambient conditions.

anaesthesia RELATED: paraesthesia, analgesia ♦ The absence of sensation. This may be general
anaesthesia, in which case the subject is unconscious, or local anaesthesia affecting sensation from
just a part of the body. Local anaesthesia may be due to influence of an anaesthetic drug or to nerve
trauma, etc..

analgesia Absence of pain sensation.

analogue RELATED: graded ♦ 1. Continuously variable. An analogue parameter can have any

aneurysm An abnormally bulging part of a vessel, usually an artery. NB nothing to do with nerves:
derives from ana= up + eurys= wide.

anion CONVERSE: cation ♦ Negative ion. NB negative ions move towards an anode (positive
electrode) in solution: hence the seemingly confusing nomenclature.

anisotropy Having different properties in different directions.

anoxia A condition without oxygen.

antagonist CONVERSE: agonist ♦ Something having the opposite effect. E.g. a chemical
opposing the action of another chemical, or a muscle pulling in the opposite direction.

ante- CONVERSE: post- ♦ Before. E.g. antenatal= before birth.

anterior CONVERSE: posterior ♦ Near or nearer to the front.

aqueous Associated with water.


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arterial pressure The pressure within an artery, relative to ambient atmospheric pressure. This
varies during the pulse between systolic pressure (maximum) and diastolic pressure (minimum). It
also varies along the length of arteries due to the effects of hydrostatic pressure and (especially in
small or obstructed arteries) the resistance of the arteries to flow. Standard clinical measurements
are normally made in a large artery at the same height as the heart.

arteriole Narrow terminal portion of an artery, leading to the capillaries. Resistance vessel. Major
site of control of blood flow to tissues, via innervation, chemosensitivity and thermal sensitivity of
the smooth muscle of the arteriole walls. Constriction increases the resistance to flow in the
arterioles. Note that it does not directly increase the pressure within the arterioles (a common
misapprehension), since they are open at both ends. Indeed, constriction generally reduces pressure
at the distal (capillary) end of the arterioles, because blood flow through the capillaries is reduced
and the capillary pressure therefore becomes closer to venous pressure.

arteriosclerosis Loss of elasticity of arteries, usually due to ageing and atherosclerosis.

arteriovenous Relating arteries and veins. The arteriovenous pressure difference is the net driving
force that causes blood to flow through a tissue.

artery Elastic, thick walled vessels carrying blood at relatively high pressure away from the heart.
Small arteries are contractile, due to smooth muscle present in the tunica media. The elasticity is
important in accommodating the blood ejected from the heart during systole without excessive rise
in pressure.

atmospheric air Composition of fresh air varies little, apart from water content (0-7%). Oxygen is
21%, N 2 78% and CO 2 0.03% of dried air. The CO 2 content can be regarded as zero for all
practical purposes in animal physiology. The total pressure (ca. 10.1kPa, 760mmHg at sea level) is
the sum of the partial pressures of constituent gases, including water vapour pressure (= relative
humidity ♦ saturated water vapour pressure).

autonomic RELATED: parasympathetic, sympathetic ♦ Relating to the autonomic nervous system.


This innervates smooth muscle, glands and visceral organs, which are not normally under voluntary
control. Subdivided principally into the sympathetic and parasympathetic efferent systems.
Autonomic reflexes are reflexes that act through these efferent systems; their afferent pathways
may be either the same as pathways that subserve conscious perceptions (as with salivation) or they
may be different (as with baroreceptor reflexes). The afferent pathways are not distinctive in any
anatomical way, and are not usually described as 'autonomic' except by association with particular
reflex actions.

axon A nerve fibre. May be myelinated or unmyelinated. A nerve is made up of many axons,
Schwann cells (supporting cells) and a sheath.

balance A balance control on a measuring instrument (e.g. an oscilloscope amplifier) allows you to
adjust the internal circuitry of the instrument for ideal operation. To adjust the balance control of an
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oscilloscope, connect the input of the amplifier to earth and gradually increase the sensitivity from
minimum. With no input (zero volts) you should expect the recording (i.e. spot or trace height) not
to change with increasing sensitivity. If it does change, adjust the 'balance' and try again from
minimum sensitivity. Once you have done this, you should not use the balance control to make
adjustments if a recording goes off-scale: you can usually use a 'shift' or 'offset' control.

band pass RELATED: filter, high pass, low pass ♦ A type of filter that lets through a range of
frequencies and cuts both higher and lower frequencies. A high pass and a low pass filter acting in
series will achieve this, though it may also be achieved in a single circuit. A band pass filter with a
narrow pass band is equivalent to a 'resonant' filter.

base In an expression of the form y=ab, 'a' is the base and 'b' is the exponent. Correspondingly,
logarithms are defined in relation to a specific 'base'. In the expression y=ab, 'b' is the logarithm to
the base 'a' of 'y'. In chemistry a 'base' is a molecule or ion that will bind hydrogen ions.

baseline RELATED: control ♦ Period of recording before a stimulus is given. Recordings in an


experiment are usually useless unless there is some baseline recording, showing reasonably stable
measurements. A response is only worth considering if it is larger than, or at least different from,
baseline fluctuations. Sometimes it is impossible to get baseline measurements: for example, you
usually cannot make baseline measurements on patients before they get sick. Then you have to rely
on control subjects (e.g. people who aren't sick) to check that they don't show the same changes
when you start to make measurements on them.

blind RELATED: double blind ♦ An experiment in which either the experimenter or the subject is
not aware of some aspect of the experimental conditions. For example, an assay might be done
'blind', with the person carrying it out not aware of the source of individual samples. This helps to
eliminate 'bias', i.e. the possibility of preconceived ideas of the experimenter affecting the results.

block 1. In physiology and pharmacology, 'block' usually means to stop something happening (e.g.
a local anaesthetic blocks action potentials in nerve fibres), or to diminish (by any of a number of
ways) the effectiveness of a drug, e.g. curare blocks the action of acetylcholine at the nerve-muscle
junction. 2. An experiment with trials carried out 'in a block' means the trials are close in time, or
not interspersed with other trials under different conditions.

boundary conditions RELATED: differential equations ♦ The conditions that determine the
particular form of solution that a differential equation will have. Sometimes these are conditions at
a physical boundary, e.g. the edge of a piece of tissue. Sometimes they are conditions at a particular
time or place, as when there is a sudden local disturbance. For example, the diffusion equation
applied to a region of tissue has different solutions depending on whether the surface is permeable
and washed by a solution free of the diffusing substance (c=0 at the surface), or is impermeable to
the substance (implying zero flux and concentration gradient perpendicular to the surface).

buffer RELATED: pH ♦ A buffer mechanism makes the effects of a disturbance less than they
would otherwise be. Particular mixtures of chemicals can buffer the concentration of an ion, for
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example H+ or Ca++. A buffer for pH is a mixture of the protonated and unprotonated forms of a
weak acid (e.g. H 2 PO 4 - and HPO 4 --. Addition of acid or base causes conversion between these two
forms, binding (or releasing) hydrogen ions. This keeps the pH relatively constant. Buffers are
most effective, for a given total concentration, when the pH is approximately equal to the pK of the
reaction: the two forms of the buffer are then in equal (and therefore quite high) concentration.

calorie Unit of energy, particularly heat. 1 calorie= 4.18 Joules. The 'Calorie' of nutrition, in a non-
scientific context, is unfortunately usually 103 calories. However, 'kcal' on a food packet will
correctly mean 103 cal, not 106 cal. Best to avoid this unit (not strictly part of S-I units) when
possible, and use Joules.

cannula RELATED: catheter ♦ A tube inserted into a vessel (usually a blood vessel) through its
wall, so that solution can be put into or withdrawn from the vessel.

capacitance Physiological 'capacitance vessels' in the circulation are ones that can contain an extra
volume of blood with little or no increase in pressure (mainly the veins, which are like floppy
tubes). Electrical 'capacitance' is related: electrical charge is like an amount of electrical substance,
and voltage is like pressure. The capacitance between places A and B is the amount of charge you
can shift from A to B per unit of voltage change between A and B. A large capacitance (measured
in farads: = Coulombs/Volt) means you need a lot of charge to get much of a voltage shift.
Myelinated nerve fibres conduct faster, largely because the myelin reduces the capacitance of the
axon.

cardiac Relating to the heart, or (as in 'cardiac sphincter') to the top end of the stomach.

carrier RELATED: channel ♦ In the context of membrane transport, a protein that binds a specific
substance and by changing conformation transports the substance to the other side of the
membrane. A carrier is very much like an enzyme. Carriers are involved in facilitated diffusion, in
which the net flux is always down an electrochemical gradient, and also in active transport. Some
carriers move two substrates at once.

catabolism Metabolism involving the destruction or disappearance of tissue. Fat stores, and
ultimately muscle tissue, are catabolised in starvation.

catheter RELATED: cannula ♦ Tube inserted into a narrow opening so that fluid can be removed
or introduced. You might sometimes insert a catheter through a cannula.

cerebral Relating to the brain.

cervical Relating to the neck, or the neck of the uterus (womb).

channel RELATED: permeability, carrier ♦ In membrane transport, a protein that spans the cell
membrane and allows substances (usually ions) to move across the membrane. A conformational
change in the protein is not involved for each ion that moves, unlike with a carrier. Much higher
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fluxes can occur than with carriers, and channels are mainly responsible for the permeability of
most membranes. Many channels are gated, i.e. can be opened or closed, e.g. as a result of binding
of transmitters, hormones or intracellular messengers, or as a result of changes of membrane
potential. Different channels also have different selectivities for the ions they let through. Single
isolated ion channels can be studied with 'patch' techniques.

chronic RELATED: tonic CONVERSE: acute ♦ Continuous, or existing for a long time.

concentration ABBREV: RELATED: activity ♦ The quantity of a solute per unit volume. It may
be measured in g/l, mols/l, kg/m3, g/100ml, etc., always with dimensions mass/volume.
Physiological concentrations are most often expressed in mols/l (i.e. the 'molar concentration' or
'molarity'). The unit of molar concentration is 1mol/l, also written as 1M. Note that it is quite
wrong to express a concentration either as 1mol, or 1M/l. These are NOT units of concentration!
Occasionally concentrations are expressed as %, or parts per million (ppm), or g/kg. These ratios
usually mean the weight of solute per unit weight of solution, or (for a gas) the volume fraction
within a mixture. '%' may also mean g/100ml.

constriction CONVERSE: dilation ♦ Narrowing of a vessel. E.g. vasoconstriction = narrowing of


blood vessels, usually due to contraction of smooth muscle.

contra- RELATED: anti- ♦ Against, opposite

contraction CONVERSE: relaxation ♦ State of activity of a muscle in which it may produce force
and/or shorten, depending on the mechanical constraints on it. Note that 'contraction' in ordinary
English means 'getting smaller'. In muscle physiology the word 'shortening' is reserved for this, and
'contraction' may occur even while a muscle is lengthening. Note also that 'activation' in muscle
physiology is also different from contraction, referring specifically to the electrical activation of the
membrane.

control RELATED: control system, control experiment ♦ Many physiological systems are 'control'

cortex The skin, or outer part of an organ (e.g. the cerebral cortex or adrenal cortex).

cranial Relating to the skull (cranium).

CT scan RELATED: CAT ♦ Computed tomography, or computed axial tomography. An X-ray


technique in which an image of the internal structure of tissue is reconstructed, not just a shadow
picture as in an ordinary X-ray.

data The results of an experiment or investigation. Strictly, it is the plural of 'datum', and you
should say for example 'These data show...'. In practice, the word 'datum' is hardly ever used and it
is probably more common to say 'This data shows...'.

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dendrite A branching, tree-like structure. Most commonly, the part of a neuron that receives
synaptic contacts.

depression RELATED: inhibition CONVERSE: potentiation, facilitation ♦ A decrease in the


activity of tissue or the amplitude of a response. If the decrease is due to the involvement of
structures (e.g. particular synapses) that appear to have the specific function of causing the
depression, then it is usually called 'inhibition'. 'Spreading depression' is a disturbance that causes a
profound but transient depression of the excitability of nerve cells in local regions of the central
nervous system. 'Clinical depression' is a mental condition characterized by extreme sadness, a
component of several clinical syndromes.

diastole RELATED: relaxation, filling CONVERSE: systole ♦ The part of the cardiac cycle during
which the ventricular myocardium relaxes, pressure falls and the chamber fills with blood.
'Diastolic pressure' is the arterial pressure at the end of diastole, i.e. the lowest arterial pressure
during the cycle.

diffusion The process by which molecules get from one place to another (or across a membrane)
by random thermal motion. 'Facilitated' diffusion is where a special molecular mechanism in a
membrane (a 'carrier') allows diffusion of certain molecules that would not otherwise get through:
rather like a parent who lifts children over a fence.

diffusion coefficient The constant that appears in either of Fick's two laws of diffusion. It has
dimensions L2T-1. Typical values for small molecules and ions in free solution are of order 10-9m2s-
1
. Typical values for diffusion of gaseous molecules are roughly 104 times larger; hence diffusion
can be an important physiological mechanism in the gas phase (for example for fluxes of O 2 and
CO 2 ) over much larger distances than in solution.

diffusion equation The partial differential equation governing the diffusion of substances. Fick's
second law of diffusion. It states that dc/dt=D(d2c/dx2+d2c/dy2+d2z/dz2), where c is concentration
and D is the diffusion coefficient. In vector notation it is dc/dt=D∇2c. It has the same form as the
equation for conduction of heat, so solutions of the diffusion equation with particular boundary
conditions (see e.g. Crank: Diffusion) can often be obtained by reference to the equivalent
solutions in a textbook of heat theory (Carslaw & Jaeger: Conduction of Heat in Solids). Solutions
are often gaussian or error functions.

dilatation, dilation CONVERSE: constriction ♦ Opening up of a hollow or tubular structure, such


as a blood vessel or the pupil of the eye. The form 'dilation' is apparently incorrectly formed from
Latin, but seems to be rising in popularity (there is no distinction in meaning).

dimension RELATED: unit ♦ The relation between the fundamental physical quantities that
correspond to a parameter. E.g. velocity has dimension length/time; pressure has dimension
force/area, or mass/(length x time2). Quantities with particular dimensions may be measured in
different units (e.g. velocity in m/s or km/hr). The units must themselves have the correct

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dimensions however. In order to compare the size of two parameters, they must have both the same
dimensions and the same units.

discrete Separate, unmixed (e.g. discrete compartments in the body). Taking one of certain specific
values (e.g. discrete frequencies at which a piano string will resonate: the fundamental and its
harmonics).

dorsal CONVERSE: ventral ♦ Relating to the back.

dose-response curve A graph of the response to an applied drug, plotted against drug
concentration or dosage.

double blind RELATED: blind ♦ An experimenter in which both subject and experimenter are
unaware of the conditions relating to each individual subjects. For example, neither patient nor
doctor may know whether the patient is taking a drug being tested or a 'placebo'. Information about
which condition relates to each subject is kept separately and used eventually at the end of the
study to analyse the results.

drift RELATED: trend ♦ A gradual, continuous change in a parameter. Often this is an unwanted
change due to an instability in a physiological system or a gradual change in the properties of a
measuring instrument (e.g. 'baseline drift').

drive RELATED: trigger ♦ A structure is said to 'drive' another if events in the first trigger events
in the second. For example, the pacemaker region drives the cardiac cycle in other parts of the
heart. 'Drive', as a noun, usually relates to a state of an animal in which its behaviour is directed to
satisfying a particular need, such as hunger, sex, etc..

drug A chemical that affects biological tissues. Often the usage is restricted to chemicals that are
used clinically or for research purposes, or ones that are abused socially. Nutrients, toxins, and
chemicals one is particularly fond of, tend not to be called 'drugs'.

-ectomy SUF RELATED: -tomy ♦ Cutting and removing. E.g. lobectomy= removing a lobe of an
organ.

electrocardiogram ABBREV: EKG or ECG) ♦ Voltage changes recorded with electrodes on the
skin, due to the electrical events of the cardiac cycle. Standard electrodes are placed on the left (L)
and right (R) arms and on the left leg or foot (L). The right leg is earthed. Additional (precordial)
electrodes may be used on the front of the chest. Three standard connections of leads are usually
made, to record the differences in voltages: I=L-R, II=F-R, III=F-L. Alternative 'augmented'
configurations are also sometimes used (aVR, aVL, aVR). The principal components of the EKG
signal are the P wave (due to atrial depolarization), the QRS complex (ventricular depolarization)
and T wave (ventricular repolarization).

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electroencephalogram ABBREV: EEG) ♦ Voltage changes due to the brain, recorded from
electrodes on the scalp.

electrolyte A salt that dissociates into ions especially when dissolved in water. An electrolyte may
be strong e.g. NaCl or weak e.g. an amino acid. Any electrolyte in solution will conduct electricity.
All tissue fluids contain electrolytes. Strong solutions of strong electrolytes (e.g. 3M NaCl) have a
high conductivity (i.e. low resistivity).

electromyogram ABBREV: EMG) ♦ Voltage changes due to muscle, recorded from electrodes on
the skin over the muscle.

electro-oculogram ABBREV: EOG) ♦ Voltage changes due to movements of the eyes, recorded
from electrodes above and below, or on either side of, the eyes.

embolism RELATED: air embolism ♦ Blockage of an artery by an embolus. Emboli in the venous
circulation generally lodge in the lungs (pulmonary embolism). Arterial (systemic) emboli can be
more serious, sometimes producing stroke, myocardial infarction or gangrene depending on the site
of obstruction.

embolus Abnormal fragments of material carried in the circulation, e.g. a blood clot (thrombus),
fat, air (as a bubble) or foreign bodies.

endo- CONVERSE: epi-, exo- ♦ Within. E.g. endogenous= arising within a tissue.

enzyme A protein that catalyses (enhances the rate of) a chemical reaction. Its action involves
binding of the substrate or substrates (reactants) and a conformational change in the enzyme itself.

epinephrine See adrenaline.

excise RELATED: dissect ♦ To cut out and remove.

exsanguinate To let out much or all of the blood from an animal.

extra- CONVERSE: intra- ♦ Outside. E.g. extravasation= something getting out of blood vessels.

fatigue A reduction in a response as a result of repeated or prolonged stimulation or activity. In the


context of muscle function, it usually means a reduction in the amount of force that the muscle can
produce.

feedback RELATED: control ♦ Information sent to an earlier stage in a multistep process.


'Negative feedback' has effects that reduce the subsequent feedback signal, or reduce its rate of
increase. This usually has a control function, keeping a parameter in the system constant. For
example, the activity of the heart raises the arterial blood pressure. If the pressure rises above a
controlled level, signals are sent via baroreceptors and the brain back to the heart, leading to a
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decrease in heart rate. 'Positive feedback' produces changes that lead to a further increase in the
feedback signal. This can lead to runaway changes in the system, and usually to an all-or-none
response (e.g. an action potential).

fibrillation RELATED: atrial flutter ♦ Repetitive high frequency spontaneous activation or


contraction of a muscle, particularly cardiac muscle. Quivering. Atrial fibrillation is faster than
atrial flutter (>300/min) and more irregular. Ventricular fibrillation is similar, and arises frequently
from electrocution or a myocardial infarct: it is normally lethal within a few minutes if not arrested
(e.g. by electric shock from a defibrillator) or unless blood flow is sustained by cardiac massage.

glyc- Relating to glucose or glycogen.

gustatory Relating to taste.

g-i tract Gastrointestinal tract, from the mouth to the anus.

habituation

Reduction of a reflex response on successive repetitions of a stimulus.

haemostasis Blood clotting.

half life ABBREV: t 1/2 RELATED: time constant ♦ The length of time for something (most
commonly a rate of radioactive decay) to fall to 50% of its current value. If the half life is constant
whatever the current value, then the decline is exponential. The half life is then ln(2) (=0.69) times
the exponential time constant for the decline.

hepato- Related to the liver.

homeo- RELATED: = homo-, iso- ♦ Same. E.g. homeostasis.

homo- RELATED: = homeo-, iso- CONVERSE: hetero- ♦ Same. Equivalent to homeo-, in words
that come from Latin instead of Greek. E.g. homogeneous (homogenous in USA) = having a
constant property everywhere.

homogeneous RELATED: isotropic ♦ Having the same properties at every place. Distinguish
carefully from 'isotropic'. For example, the structure of contractile proteins within a skeletal muscle
cell is homogeneous, but not isotropic. Any place in the cell has similar characteristics to any other,
but these characteristics are very different in different directions.

hormone RELATED: endocrine ♦ A chemical that is released from cells and transported to its
target organ in the blood.

humoral Relating to body fluids.


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hypothesis RELATED: null hypothesis, theory ♦ A suggestion; part of a theory; a postulate. Many
of the explanations in physiological textbooks for how things work are really hypotheses. They
may be widely accepted, or sometimes rather uncertain and controversial. It is seldom possible in
science to prove that a hypothesis is correct. You should always treat scientific explanations with a
healthy scepticism, regarding them as in principle capable of being proved wrong or requiring
some modification in the light of new experiments. If you are asked for a hypothesis to explain
some data, it is always a good idea to think of 2 hypotheses: you may then be able to devise a
crucial experiment to distinguish them.

in vitro RELATED: culture CONVERSE: in vivo ♦ Literally, in glass. Often refers to a procedure
carried out on cells or tissue isolated from the body and maintained in a tissue bath.

indication A sign, hint, or suggestion. In clinical terminology: a circumstance that suggests that a
particular therapeutic regime or diagnostic test is merited (as opposed to a 'contra-indication': a
circumstance that suggests that a course of action might be unwise).

infarct A non-functioning, or totally dead, region of tissue (often resulting from ischaemia).

infinitesimal RELATED: differential ♦ The limit of very small quantities, analogous to the
reciprocal of 'infinity'.In one sense, an infinitesimal quantity is simply zero. However, you may be
interested in the ratio of two things as they become infinitesimal, when it is not helpful to think of
them as zero.

inhibition RELATED: depression CONVERSE: excitation, facilitation ♦ An influence that reduces


activity in a tissue or reduces the response produced by a stimulus. Synaptic inhibition can operate
either by hyperpolarizing a cell (subtracting from the effects of excitation) or by reducing the
amount of depolarization caused by excitation (diminishing the excitatory influence, with no effect
unless excitation is taking place). The second situation can arise with 'presynaptic' inhibition, and
also as a result of a postsynaptic conductance increase that opens channels that tend to keep the
membrane potential close to the resting potential. If the decrease of a response outlasts the
influence that brings it about, it is usually called 'depression'.

innervate One tissue is innervated by another if it contains terminals of axons arising from cell
bodies in the second tissue. These might be synaptic terminals or sensory terminals (as in the
sensory innervation of the skin from cells in the dorsal column nuclei). The term is essentially
anatomical, and does not relate to the processes of activation or synaptic transmission.

input CONVERSE: output ♦ An influence or a physical route into a structure, that is capable of
affecting it. The influence may be the physical entry of something (e.g. the input of food into the
stomach) or the passage of information (e.g. action potentials in the innervation of the stomach).
Cables connecting pieces of equipment usually have clearly defined input and output ends: signals
or electric power are usually conveyed in one specific direction, which you should identify to make
sense of the wiring.

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interstitial RELATED: extracellular ♦ Relating to the interstices, or spaces in between things
(usually cells). The interstitial space is the extracellular compartment.

intra- RELATED: inter- CONVERSE: extra- ♦ Inside. 'Intracellular' means inside cells, quite
different from 'intercellular'.

invasive CONVERSE: non-invasive ♦ An 'invasive' procedure involves putting something into the
tissue under study, removing a sample of it ('biopsy'), or performing some surgery. Radiation, X-
rays, etc. are usually regarded as 'invasive' because the invisible penetration by particles or waves
is capable of doing damage.

inverse Two things are inversely related if one goes up when the other goes down. For example,
pH and hydrogen ion concentration are inversely related. If the relation is not strict (as it is in this
case, where there is an equation that always relates the two things exactly), then it is more usual to
talk about a 'negative correlation' than an 'inverse relationship'.

ischaemia RELATED: hypoxia, anoxia ♦ Cessation of blood flow. This is a normal condition in
some tissues, for example in many muscles during maximal voluntary contractions. If continued for
too long, the products of metabolism build up, substrates become depleted, and temporary or
irreversible loss of function (infarction) may result.

iso- RELATED: hypo-, hyper- ♦ Having the same value, or property. E.g. isotonic, isomer

isolated RELATED: in vitro ♦ A tissue separated from its normal inputs. It might be kept in the
body for study ('in situ') or it might be studied 'in vitro'. An 'isolated stimulator' is an electrical
stimulator in which neither of the output terminals is connected to earth; this is useful for reducing
the current during a stimulus that may flow through a recording device, causing a 'stimulus
artefact'.

i-m Intramuscular (usually referring to an injection site).

i-p Intraperitoneal. An injection given into the abdominal cavity (belly).

iso- equal, same, uniform

isotonic RELATED: isosmolar ♦ (1) Having the same tonicity, or the same osmotic effect on cells
as the fluid in their normal environment. This is not necessarily the same as having the same
osmolarity (see tonicity) because some solutes contributing to the osmolarity of a solution may be
permeant and equilibrate across the membrane, and so have no effect on cell volume, or only a
transient effect. (2) In muscle physiology, an isotonic contraction is one during which the tension
is constant (contrast isometric) .

-itis SUF Inflammation of.

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i-v Intravenous (an injection or infusion into a vein).

latency A time delay between a stimulus and the beginning of a response. Don't use the term
'latency' when you are talking about the time to the peak of a response. The word literally means
'hidden' time: the time after the stimulus when there is still no sign that any response is going to
occur.

lesion RELATED: trauma ♦ A region of injury. The verb 'to lesion' is sometimes used for the
deliberate induction of tissue damage for experimental purposes.

ligand A chemical or ion that binds to another chemical.

ligate RELATED: ligature ♦ To tie closed, for example, a blood vessel or duct from a gland.

lyse To break, dissolve or destroy. Often refers to cells bursting, for example, due to a hypotonic
extracellular solution.

macro- CONVERSE: micro- ♦ Large. E.g. macrophage= a large cell that engulfs particles. In
computer jargon, a 'macro' is a single command that initiates a complex set of commands.

macroscopic CONVERSE: microscopic ♦ Relating to large-scale features of something

magnetic resonance ABBREV: MRS, MRI RELATED: NMR ♦ Magnetic resonance. A technique
for studying the internal structure and chemistry of tissue by observing its properties in a high
magnetic field. MR spectroscopy (MRS) gives information about the different chemicals present.
Imaging techniques (MRI) give images of the structures that differ in their chemistry, which can
reveal tumours, dead tissue, etc.

malignant RELATED: cancer CONVERSE: benign ♦ A condition that gets worse if not treated.
Particularly refers to a tumour that invades and destroys other tissues.

median RELATED: quartile, percentile ♦ The value within a distribution that is exceeded by half
of the data points. The 50% percentile.

medium RELATED: culture ♦ Nutrient fluid suitable for growth or maintenance of cells or tissue
in vitro.

mega- ABBREV: M RELATED: SI units ♦ One million. 10^6 or 106.

megaly SUF Large. E.g. megalomania = delusion of grandeur.

metabolism The chemical reactions that occur in the body. Chemicals ingested or manufactured in
the body are either metabolised, excreted or accumulated.

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metabolite A product of metabolism of nutrients or of a particular specified substance.

micturition Urination.

milli- ABBREV: m RELATED: SI units ♦ One thousandth. 10-3. For example, the commonest unit
of voltage used in physiology is the millivolt (mV). Note that the abbreviation (m) is the same as
that for a metre. This does not lead to confusion if you use units correctly. If 'milli-' is used, it
always precedes a symbol for an ordinary unit, and never stands on its own. Separate units that are
multiplied together should always be separated by a dot (period). Thus 1ms = 1 millisecond; 1m.s =
1 meter second.

mmHg The unit of pressure most commonly used for measuring blood pressure. 1mmHg is
approximately 133 Pa (pascals). The unit is retained, despite not being an SI unit, because the
commonest and most accurate instrument for measuring blood pressure is a mercury
sphygmomanometer, which gives a direct reading in mmHg.

mode RELATED: mean ♦ The most common, or most probable value in a distribution. Seldom a
useful concept in physiology, though it is often about the same as the mean or median.

morbid Diseased. Distinguish 'morbidity' (the extent to which something, for example an infection,
causes disease) from 'mortality' (the extent to which it causes death).

motility Movement, particularly of cells or parts of cells.

nephro- RELATED: renal ♦ Relating to kidney. E.g. nephritis= inflammation of kidney.

non-invasive CONVERSE: invasive ♦ A technique that doesn't require entry into the tissue being
studied, or damage to it. Many recent medical advances have arisen through the development of
non-invasive techniques that assist diagnosis (e.g. Magnetic resonance).

noxious A stimulus that is painful or unpleasant.

observation A measurement, or observation of a qualitative feature. Observations of the responses


to skin trauma might include changes of skin colour, swelling, changes of heart rate, vocalisation,
etc.

occlusion A blockage in for example, a blood vessel. The response to one stimulus is sometimes
said to 'occlude' the response to another if the response to both stimuli presented together is less
than the sum of the two separately. This occurs in the nervous system, for example, where each
stimulus induces an all-or-none response in some of the same neurons.

oedema RELATED: = edema (US) ♦ Swelling

olfactory Relating to smell.


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ortho- Straight, normal. E.g. orthodontics = straightening of teeth.

output RELATED: projection CONVERSE: input ♦ A structure leading from a system or


something (a chemical, signal or a form of energy) produced by the system.

overshoot RELATED: undershoot, transient ♦ Part of a response to a continuing stimulus, in which


the response transiently exceeds the level at which it finally settles. Note that it is not really the
converse of an undershoot.

para- Alongside, resembling. E.g. E.g. parathyroid= gland beside the thyroid, paraesthesia=
abnormal (but not absent) sensation, paramedical= alongside medicine. Can also mean a defence
against something (e.g. parasol), and in chemistry two positions opposite each other on a molecule.

partial pressure in a gas mixture RELATED: vapour pressure ♦ The portion of the total pressure
of a gas mixture that is due to a particular constituent. The total pressure is the sum of the partial
pressures, and the proportions are the same as the proportions of the quantities of gas, either by
numbers of molecules or moles, or by volume (measured at a fixed pressure). For example, normal
expired air contains (after drying) about 5% CO 2 by volume (5% of the molecules are CO 2 ). The
CO 2 partial pressure (P CO2 ) is therefore about 0.05 of an atmosphere (38 mmHg or 5.0 kPa). Since
part of the pressure in the alveoli (about 47 mmHg or 6.3 kPa) is water vapour pressure the CO 2
partial pressure in the alveoli would be less: 0.05 of (760-47) mmHg = 36 mmHg, or 0.05 of (101-
6.3) kPa = 4.7 kPa

partial pressure in a solution RELATED: partial pressure in a gas mixture ♦ The partial pressure
of a gas (like O 2 or CO 2 ) in a solution is the partial pressure in a gas mixture in equilibrium with
the solution. It is not directly related to how much of the gas is in the solution. Under normal
conditions, arterial blood has P O2 = 100 mmHg and P CO2 = 40 mmHg, close to the values in the
alveolar gas mixture with which it has equilibrated. At these normal partial pressures, the amount
of O 2 and CO 2 in the blood may vary depending on how many red cells and much haemoglobin the
blood contains.

pathogen An organism or chemical that causes disease.

pathological Diseased, disordered, or abnormal.

pathophysiology Study of the function of tissue under pathological conditions.

perception Awareness of something, especially a sensation due to a sensory input. There are many
sensory inputs that do not lead to conscious perceptions (e.g. afferents from arterial baroreceptors
and chemoreceptors). Curiously, subjects can deny any perception, or any knowledge at all of a
stimulus, yet be able to identify accurately some of its properties when forced to guess what they
are (e.g. in the phenomenon of 'blindsight' in patients who have total lesions of the primary visual
cortex).

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perfuse RELATED: superfuse, infuse ♦ To pass fluid through. E.g. an isolated salivary gland
might be perfused with saline.

peri- RELATED: epi- ♦ Surrounding, around. E.g. perineurium= sheath around nerve, perinatal=
around the time of birth.

perinatal Around the time of birth.

permeability RELATED: diffusion, transport ♦ Passage of a substance by diffusion across a


membrane. A membrane is said to be 'permeable' to a substance; the substance is said to be
'permeant'. The permeability of a membrane only allows a 'passive' net flux to take place, i.e. in the
direction down the electrochemical gradient of the permeant substance. If net movement occurs in
the other direction, the substance must be subject to 'active transport'. Quantitatively, the
'permeability coefficient' for an uncharged substance, or for an ion under conditions when the
membrane potential is zero, is the flux per unit area, divided by the concentration difference
between the two sides of the membrane (dimensions: LT-1).

pH RELATED: pCa ♦ A measure of acidity. pH = - log 10 (molarH+ concentration). Note (a) that it is
a logarithmic measure, so a change of 1 unit in pH corresponds to a 10-fold change in H+
concentration, and 0.3 units to a 2-fold change in concentration. (b) it is an inverse measure: low
pH corresponds to high H+ concentration and high acidity. It is easy to remember that pH=7
corresponds to 10-7 M and pH=8 to 10-8 M. Normal plasma pH, ca. 7.4, is of course in between.

phage SUF Eating, engulfing. E.g. phagocyte.

phase RELATED: time course, amplitude ♦ 1. One of the parts of the time course of an event. For
example, the rising or falling phase of an action potential; the recovery phase after exercise. 2. For
a sinusoidal function: a parameter that indicates where you are on the waveform, relative to when
the waveform crosses the axis in a positive-going direction (phase=0). This is measured as an
angle: 2π´t/T radians or 360×t/T degrees, where t is the time relative to the crossover and T is the
period. A 'phase shift' is the amount you have to shift one sinusoidal waveform relative to another
(of the same frequency) to make them have the same crossovers.

photomultiplier tube An extremely sensitive light sensing device that transduces a weak light
signal into an electrical signal.

physiological CONVERSE: unphysiological ♦ Characteristic of the normal workings of the tissue.


If an experimental condition or a type of stimulation is said to be 'physiological', this means that it
is within the range of conditions normally experienced by the tissue within the animal. Experiments
under 'unphysiological' conditions are often enormously helpful in understanding how the tissue
works. For example, study of the force produced by muscle cells when stretched farther than the
skeleton will normally permit them to be stretched (i.e. beyond their 'physiological range') were
instrumental in establishing the sliding filament theory of muscle contraction.

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placebo An inactive chemical given, as a control, to a subject in an experiment. New drugs cannot
be claimed to be effective unless they are better than a placebo in a controlled, preferably 'blind',
study. Many clinical conditions show a surprising improvement in many subjects following
administration of a placebo (a 'placebo effect'). It has been suggested that the size of a placebo
effect is a measure of the social and psychological skills of a doctor, since undoubtedly these play a
part in therapy, and with a placebo there is nothing else to help.

plasma RELATED: serum ♦ The extracellular fluid of the blood, in which the cells are suspended.
If blood is prevented from clotting, the cells will settle out leaving the relatively clear, yellowish
plasma.

prognosis The predicted course of a disease or disorder.

prophylactic A drug or a preventative measure taken to avoid disease or infection.

proteinuria Presence of protein in urine, sometimes indicative of kidney or heart disorders.

protocol The plan for an experiment.

pylorus The narrow end of the stomach (pylorus) leading to the duodenum.

refractory period A period in which a cell or tissue is unresponsive to stimuli (absolute refractory
period) or has a raised threshold (relative refractory period) following a preceding period of
activity.

regulate RELATED: control ♦ To control, in the sense of keeping something constant (e.g. body
core temperature) or constantly adjusted to suit current requirements (e.g. fluid excretion in the
kidney).

renal Relating to the kidney.

resistance vessels The arterioles and small arteries, which provide the greatest resistance to blood
flow of all the vessels through which the blood flows in sequence through a tissue. Consequently,
the biggest drop of pressure is along the resistance vessels.

resolution 1. The degree of detail contained in a visual image (e.g. expressed by the number of
pixels). 2. The acuity or resolving power of a sensory system or an instrument. 3. The termination
of an acute phase of a disease, especially a stage at which inflammation disappears.

resolution of measurement RELATED: accuracy ♦ Ability of a measurement to distinguish


slightly different conditions. Resolution may be limited by the size of incremental steps in the
measurement (e.g. a digital pH meter may give readings to 2, 3, or 4 decimal places) or by
inconsistencies in behaviour. It may be possible to identify the sources of some inconsistencies and
reduce them (e.g. by placing the instrument in a standard state immediately prior to a
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measurement). Limitations due to fluctuations that are truly random can usually be improved by a
factor SQRTn by averaging n independent measurements. Resolution is just one of the factors that
limits the accuracy of a measurement.

respiration (1) The act of breathing. (2) metabolic reactions involving oxygen as a reactant.

retrograde Moving backwards or in an abnormal direction or (as in retrograde axonal transport) in


the opposite direction to something else (the action potentials).

Ringer solution RELATED: Krebs solution, ♦ A simple saline solution that is sufficiently similar
to the normal environment of a tissue that the tissue continues to behave fairly normally when
immersed in it. Named after Sidney Ringer, who discovered that calcium ions are an essential
component of the physiological environment of cardiac cells. Appropriate Ringer solutions vary for
different tissues and different species, and according to the needs of a particular experiment. Ringer
solutions are usually not bubbled with a gas mixture containing CO 2 , and therefore normally have
a lower than normal bicarbonate concentration to ensure a physiological pH.

saline RELATED: Ringer solution ♦ A solution of salts, usually one suitable for bathing or
injecting into tissues. The simplest saline for clinical and mammalian use is 0.9% NaCl (approx.
150mM), which is roughly isosmotic with cells. More elaborate solutions for maintaining tissues in
good physiological condition are often named after scientists who worked out their formulae, e.g.
Ringer, Krebs.

serum RELATED: plasma ♦ The fluid that separates from blood when it clots. Approximately
equivalent to the plasma without much of its dissolved proteins.

sigmoid A curved graph that always rises as you move to the right, but for which the gradient
increases at first and then decreases. If you sketched a graph of the gradient, this would rise and
then fall. This is a common shape of graph in biology, for example for dose-response curves,
stimulus-response curves and for chemical reactions that involve cooperative phenomena, such as
the oxygen dissociation curve for haemoglobin. The word means S-shaped, but it's really more like
the shape of an integral sign.

supra- RELATED: super-, hyper- CONVERSE: sub- ♦ Above.

supramaximal RELATED: maximal, threshold ♦ Above maximal. A supramaximal stimulus is


more than big enough to have the full effect: increasing it further will not increase the response.
The stimulus is therefore on the plateau of a stimulus- response graph. This may be because the
stimulus is above the threshold for all of the cells contributing to a response.

suture A stitch made with needle and thread or a metal clamp, to hold tissue together.

sweep The left to right movement of the spot on an oscilloscope.

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symptom RELATED: sign ♦ A change noticed by a patient, that indicates an underlying disorder.
A 'presenting symptom' is one that leads the patient to seek medical advice: it is not necessarily
indicative of the most serious aspects of the disorder.

syncytium A set of cells that are coupled in some way, so that they behave in certain respects as if
their cytoplasm was continuous. For example, electric current can pass through the gap junctions
between cells in the heart, allowing an action potential to propagate throughout the heart without
synaptic transmission.

syndrome A set of symptoms and/or signs that are characteristic of a particular disorder.

systole RELATED: diastole ♦ The time of contraction of the ventricles of the heart, when arterial
blood is expelled into the aorta. Systolic pressure is the maximum arterial pressure reaches during
the cycle, at the end of systole.

temporal 1. Relating to time. 2. Relating to the temple, or side of the head.

tetanus RELATED: twitch ♦ In muscle physiology, a continuous contraction produced by stimuli


or action potentials at a high frequency. A 'fused' tetanus is one in which the frequency is high
enough that responses to individual stimuli cannot be distinguished. Tetanus (or lockjaw) is also a
disease in which sustained muscular contractions occur. The word is used in neurophysiology to
denote high frequency stimulation, though this is often considered improper since the origin of the
word has to do with the tautness of muscle, not with the frequency of stimulation.

threshold RELATED: excitability ♦ Minimum strength of stimulus that elicits a response.

-tomy RELATED: -ectomy ♦ Cutting. E.g. lobotomy= incising a lobe of an organ.

tonic CONVERSE: phasic ♦ Continuous. A tonic response is one that is maintained during the
period of a stimulus, i.e. that does not show complete adaptation. Muscle 'tone' is a continuous
level of contraction, arising usually from tonic activity in the innervating nerves (e.g. in skeletal
muscles and vascular smooth muscles).

tonicity RELATED: osmolarity ♦ In physical chemistry, this is the same as 'osmolarity'. In


physiology, it relates specifically to the behaviour of cells in a solution. It is a measure of the
tendency of a solution to make cells swell or shrink. An 'isotonic' solution leaves cells at normal
volume. A hypotonic solution (e.g. a more dilute solution, with lower osmolarity) makes cells
swell; a hypertonic solution makes them shrink. Substances in solution that readily pass through
membranes (e.g. urea) contribute to the osmolarity of a solution, but not to its tonicity. A 1M urea
solution has a higher osmolarity than blood plasma, but is hypotonic. Water will enter the cells
along with the urea, and make the cells burst.

toxin A poisonous or harmful substance. Many useful drugs have toxic effects.

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transport In the context of membrane physiology, (verb) to move across a membrane.

trauma RELATED: lesion ♦ A painful, harmful, or destructive event, or the wound caused by such
an event. The word is used in relation to both physical and psychological damage.

trend RELATED: regression, correlation ♦ 1. An approximate relation, or a generalisation that is

trophic Relating to nutrition or growth. A trophic influence is one that influences growth or
maintenance of a tissue.

tropic SUF Related to direction. Turning. E.g. trophotropic= turning or moving towards food.

-uresis Relating to, or appearing in, the urine. E.g. diuresis= extra urine, proteinuresis= protein in
the urine.

vaso- Relating to a vessel (i.e. in anatomy, a tube). Usually relating to a blood vessel (e.g.
vasoactive= affecting blood vessels), but NB vasectomy= removal of a part of the vas deferens.

vector RELATED: scalar ♦ A quantity that has direction as well as magnitude. You can represent it
visually as an arrow. For example, the velocity of blood flow is a vector, having magnitude and
direction everywhere within a blood vessel. If the flow is laminar in a straight vessel, the direction
is everywhere the same, but the magnitude falls off towards the vessel walls. At a branch point, the
vectors in different places have different directions. If there is turbulent flow, the vectors fluctuate
somewhat chaotically with both position and time. You can 'resolve' a vector into components in 3
perpendicular directions. NB vectors with zero magnitude do not have a defined direction.

vectorcardiography A way of analyzing electrocardiogram signals in which the signals are


displayed as a spot moving in two or three dimensions, corresponding to the changes of both
amplitude and electrical axis of the electrocardiogram during the cycle. The spot traverses loops,
which give a direct visual indication of the electrical axis.

ventilate RELATED: breathe ♦ To cause air to flow through the airways.

vivisection Literally, means to cut something living. Refers to experiments on living (including
anaesthetised) animals. Anti-vivisectionists are those who oppose the use of animals for
experiments. Opposition to animal experiments should be distinguished from opposition to cruelty
to animals. Most people who perform animal experiments for biomedical and veterinary research
are wholly opposed to cruelty and would argue strongly that their work causes little or no suffering
to animals, and that where suffering is caused (as is inevitable, for example, in some research on
pain) it is more than balanced by the alleviation of suffering due to the successes of such research
in leading to clinical advances.

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