Undergraduate – Graduate

Male Reproductive System Histology Lecture Series

Larry Johnson, Professor

Veterinary Integrative
Biosciences
Texas A&M University
College Station, TX 77843
 Objectives

To examine the testis (which produce spermatozoa), excretory ducts

(which transport and mature spermatozoa), and accessory glands
(whose secretions support the viability of spermatozoa) for
characteristics and functions the male reproductive system.

To learn what structures and hormonal regulation facilitate the male

gonad to produce an exocrine secretion (the spermatozoon) and an
endocrine secretion (testosterone).
 Outline
History
Spermatozoon
Spermatogenesis
Sertoli and Leydig cells
Hormonal control
Epididymal and
accessory glans
characteristics and
functions
Fertile ejaculate
 First scientific endocrine
study involved the testes
of roosters studying the
observable effects of
hen testosterone on the
secondary sex structures.

Transplanted tested
maintained wattles and
comb growth in castrated
roosters.
 Functions of Male Reproductive System are to:

- produce, maintain, and transport mature spermatozoa (the

male gametes),
- produce nutritive and protective fluid (semen), and
- discharge the spermatozoa-containing semen within the
female reproductive tract during mating.
Primate
sperm
Mouse
spermatozoon
Rat Bull
Human
 Details of spermatozoa

The
annulus marks the
end of the middle
piece (houses the
mitochondria) and
beginning of the
principal piece of the
Horse spermatozoon
 Horse

Middle piece with mitochondria

 Human Human

Spermatozoa are produced

Horse in seminiferous tubules in the testis.
 Human testis

Leydig
cells

Seminiferous Slide 165

tubules
 Human testicular capsule

Vein

Artery

Mesothelium

Slide 165 34455

 Human testis toluidine blue
Seminiferous
19680 tubules

Leydig
cells
Seminiferous
epithelium

Spermatozoa are produced in the seminiferous epithelium lining

the lumen of the testicular tubules. Testosterone is synthesized
by Leydig cells located between seminiferous tubules.
 Human testis toluidine blue
Germ cells

467

Two kinds of cells present in the seminiferous

epithelium: Sertoli cells and germ cells: Sertoli
cells nurse germ cell development. Nuclei of Sertoli cells 19680
Horse Seminiferous Tubules
 Seminiferous tubules are composed of
seminiferous epithelium and boundary cells:

Myoid Cells

Sertoli Cells

Germ Cells
Spermatogonia,
Spermatocytes,
Spermatids
Human
 Seminiferous Tubules
composed of:

Myoid cells

Sertoli cells

Germ cells
Spermatogonia,
Spermatocytes,
Human
Spermatids
 Meiotic activity

UT165 larger secondary spermatocyte nuclei and smaller Golgi phase spermatid nuclei
human testis
 metaphase figures in meiosis #92 secondary spermatocytes

Pachytene primary
spermatocytes

Letotene primary
spermatocytes

To find secondary spermatocytes, one needs to find a tubule in stage VI of the spermatogenic cycle
with metaphase figures in meiosis and no (almost no) pachytene primary spermatocytes. The
pachytene primary spermatocytes are the immediate precursor to secondary spermatocytes.
 Early and late spermatids
Slide 92: Testis Leydig (interstitial)
cells

Tunica albuginea

Residual bodies
Basement membrane

Myoid cell

Sertoli cells

Seminiferous tubules
 Spermatogenesis is Divided into
3 Main Events
Event Cell Type Duration

Spermatocytogenesis Spermatogonia 27 Days

Meiosis Spermatocytes 24 Days

Spermiogenesis Spermatids 23 Days

Combined Duration = 74 Days

Spermatocytogenesis

Meiosis

Spermiogenesis
Spermatocytogenesis Has
Two Functions

Produces primary spermatocytes which result in

the production of sperm 47 days later.

Produces stem cells which insure a constant

supply of germ-cell precursors throughout life.
Spermatocytogenesis

Horse
 Meiosis (only in spermatogenesis
and oogenesis)
Exchange of genetic material in homologous
chromosomes (leptotene, zygotene,
pachytene, and diplotene steps of
development)

Produces haploid condition of gametes

 Meiosis
Exchange Of Genetic Material
Secondary
Spermatocytes
 Meiosis
Produces Haploid Condition of Gametes
 Spermiogenesis
(differentiation of spermatids with round nuclei
to those characteristic of spermatozoa)

Acrosome from Golgi

Nuclear condensation and

elongation with appearance
of the spermatid manchette

Flagellum (projects through

the flagellar cannel during
development)

Shedding excess cytoplasm Flagellar cannel

Manchette
 Golgi phase

Cap phase

Elongation phase

Maturation phase
Maturation phase
 Spermiogenesis

human Horse

Manchette
Golgi

Developing acrosome

Spermatid nucleus
 Cis face
Golgi

Black acid
phosphatase
precipitates in Acrosomal
the developing cap
Cytosol Transport vesicles acrosome

Trans face
Nucleus Golgi
Spermatid showing the developing acrosome
over its nucleus. Acid phosphatase enzymes
(black precipitates) first appear in the trans face
of the Golgi apparatus and are transferred to the Acrosomic vesicle
developing acrosome via transport vesicles. Testis of a spermatid
Manchette is a transient organelle Flagellar
as it is not found in spermatozoa cannel
Flagellar cannel
 When the annulus migrates to end
of middle piece, it removes the cell
membrane of the flagellar cannel
from the surface of the developing
flagellum in the middle piece region
and allows mitochondria access to
that portion of developing tail.
Annulus Mitochondria are found only in the
Flagellar middle piece on spermatozoa as that
cannel is the only region with access.
Residual bodies Spermatids and
Spermatozoa
Residual bodies
 Sertoli Cells
• Provide support and nutrition to developing germ cells

• Release spermatids as sperm

• Phagocytize degenerating germ cells and residual bodies

• Secrete:
– Androgen binding protein
– Calmodulin
– Plasminogen activator
– Inhibin

• Blood testis barrier

Human – infertile man
 Occluding
junctions between
Sertoli cells

Blood-testis occluding
junctions between sertoli
cells in seminiferous
tubules
 Human spermatogenesis: path followed by given cell

Basal compartment
with spermatocytogenesis Blood Testis Barrier
Adluminal compartment with meiosis and spermiogenesis
 Intercellular Bridges
(cytoplasmic bridges)
Cause - incomplete cytokinesis

Found among clusters of spermatogonia,

spermatocytes, or spermatids (never between cells
in different steps of development, e.g., never
between spermatogonia and spermatocytes)

Possible functions
Mediate both differentiation and degeneration of
spermatogonia
Maintain synchronous development
Intercellular
Bridges

Occluding junctions between Sertoli cells

are still above
but now appear below the
zygotene spermatocytes
Efficiency of Spermatogenesis

Species DSP/g (106)

--------------------------------------------
Rabbit 25
Hamster 24
Boar 23
Rat 20-24
Rhesus monkey 23
Ram 21
Stallion 16-19
Bull 12
Human 4-6
 Spermatogenic Cycle Length
Species Duration (Day)

Prairie Mole 7.2

Hamster 8.7
Mouse 8.9
Rhesus Monkey 9.5
Rabbit 10.7
Stallion 12.2
Rat 12.9
Bull 13.5
Beagle Dog 13.6
Human 16.0
Testicular interstitium
Primary
human
Sertoli cell nucleus
spermatocytes 19680

Fibroblasts,
Myoid
cells
Lipid droplets ule
n
in Sertoli cells Artery Ve

p h atic
Lym l
e
Mitotic figures vess
in dividing
spermatogonia
to produce Spermatogonia
primary
spermatocytes

Nerve , Leydig cells

 19680
Leydig
cells Myoid
cells. Capillaries

Nerve Since developing

germinal cells
require a higher
(100 fold)
Lipid droplets concentration of
testosterone than
Lymphatic do other cells,
Seminiferous vessel Leydig cells are
epithelium located close to
seminiferous
tubules.
Leydig cells
Artery
Fibroblasts
Lymphatic
vessel
 Leydig cells Leydig cells ???

Nerve

19670

Odd : What appears to be Leydig cells inside the nerve in the human testis.
Horse
 Hormonal Control of
Spermatogenesis

Hormone Cell Stimulated In Spermatogenesis

FSH Sertoli CellsSpermatocytogenesis Spermiation

LH Leydig Cells Meiosis (Testosterone)

 Deleterious Influences on
Spermatogenesis

• Heat

• Irradiation
• Chemicals

• Aging
Extragenital Components of
the
Male Reproductive System
Excretory Ducts

Composition (Rete
Testis, Efferent Ducts,
Epididymis, Ductus
Deferens, Ejaculatory
Ducts, Urethra)
19709
Human testis: junction of seminiferous tubule and rete
testis for sperm to exit (toluidine blue)

Junction of
seminiferous tubule
and rete testis tubule

Region of the
mediastinum testis

Rete
testis
467 Testis and epididymis – efferent
duct and epididymis

Profiles of the
epididymal duct

Efferent Seminiferous
ducts tubules

Rete testis
tubules
 Efferent ducts True cilia on their apical
surface help move sperm
Efferent ducts have a characteristic scalloped through the duct.
luminal profile due to alternating groups of high and
low columnar cells in the lining epithelium

Lumen

199
Horse efferent duct

19673
 True ciliated cells (efferent duct) and stereociliated cells
(epididymis, with sperm in lumen) of psudostratified columnar
epithelium (toluidine blue)

19678
Efferent duct

Epididymis
 Epididymis Head

Efferent duct

19673
Efferent duct
199 Extremely long (30µm), branching
microvilli (stereocilia) projects from
the apical surface of these cells
Stereocilia

Head
Tail of epididymis Smooth muscle
layer is thicker in
more distal
regions of
epididymis
467
19716
Human
Human
DUCTUS DEFERENS IN THE SPERMATIC CORD
Psudostratified columnar epithelium,
but lower in height

Epithelium
and its
lamina
propria,
showing
longitudinal
folds into
Blood vessels and ductus deferens the lumen

Nerve
196 Smooth muscle,
extremely thick
layers
Mechanisms of Sperm Transport
Location Force

Seminiferous tubules Bulk flow (10 ul/g/hr) minor

Contractions of myoid cells

Rete testis Bulk flow ciliary action

Efferent ducts Bulk flow ciliary action

Epididymis Contractions of smooth

muscle

Ductus deferens Contractions of smooth

muscle during ejaculation
speed is 800 mm/second
 Spermatic cord

is
y m
d
p idi
E

Testis

Spermatic cord 38

Human testis and epididymis 19678

 Epididymal
Spermatozoan
Maturation
Fertility
Motility

Nature of plasma membrane

Mitochondrial structural stability

Chromatin stability
168 Seminal vesicle, monkey

Branched network
produces a
"honeycomb"
appearance
 Prostate is a firm mass of collagenous
connective tissue and smooth muscle
that is invaded by numerous glandular
outpocketings of the urethra

Prostate Lumen
of gland

271
Note the abundance of smooth
muscle in its interstitium.
 Prostate

169
BULBOURETHRAL
GLAND
Two main function of male
reproductive system are to:

Produce male gametes

Deliver male gametes

 Penis – transitional epithelium and
surrounding spongy cavernous of penal urethra
Cavernous
erectile tissue

Nerve
Smooth
muscle

Transitional
epithelium
Spongy cavernous

Dense connective tissue bands surround the cavernous erectile

tissue. There is erectile tissue (spongy cavernous) that surrounds

277 the penile urethra. This allows the urethra to stretch when seminal
fluids are traveling down its length when the penis is rigid.
 Variations in the
Microvasculature
Common
Arteriole  Capillary 
Venule

Shunts
Arteriole  Metarteriole
 Venule

Artery  Av Shunt  Vein

Fig. Mechanism of erection
Erection occurs as a complex process, constituted
by psychological, neurological, hormonal, and
vascular factors.
the penis is composed of three basic anatomical
structures – two longitudinal cavernous bodies (a
kind of chambers) and one spongy body, including
the urethra. These are the cavernous bodies that
(supplied by respective arteries) increase their
volume during erection, owing to the inflowing blood.
arteries (as opposed to veins) have their muscular
layer composed of smooth muscles which – by
dilation or contraction – regulate the blood flow.
Sexual arousal and increasing activity of the autonomic
nervous system stimulates the release of neurotransmitters at
nerve endings in the cavernous bodies or in the endothelium
of the arteries. This leads to secretion of NO – nitric oxide,
which is one of the strongest smooth muscle relaxants. With
dilated cavernous arteries, the amount of blood flowing into
the penis increases, and its outflow is hindered by a
physiological compression of some specific veins. Moreover,
contraction of the ischiocavernous muscle stabilises the penis
in erectile position. A key factor of effective erection is the
condition of the vascular system, ensuring a proper perfusion
of the reproductive organs. Any pathologies of this system
(e.g., atherosclerosis, coronary disease, hypertension) lead to
problems with erection.
Accessory Glands - Composition and
Secretion
Accessory glands Secretion
Seminal vesicles Fructose fibrinogen

Prostate Citrate fibrinolysin

Bulbourethral gland Mucus-like lubricant

Glands of Littré Mucus

Functional Properties of the Accessory Glands

Specific contributions of seminal plasma as measured by the

split ejaculate method

Fraction of
Ejaculate Contains Source
First 90% Of All Citrate Prostate
90% Of All Sperm Ductus Deferens
Last 90% Of All Fructose Seminal Vesicles

Developmental response to androgens

CHARACTERISTICS OF FERTILE
HUMAN EJACULATES

GOOD VISCOSITY (CLOT

THEN DISPERSE)
about 3 ml in volume
 Many illustrations in these VIBS Histology YouTube videos were modified
from the following books and sources: Many thanks to original sources!

Bruce Alberts, et al. 1983. Molecular Biology of the Cell. Garland Publishing, Inc., New York, NY.
Bruce Alberts, et al. 1994. Molecular Biology of the Cell. Garland Publishing, Inc., New York, NY.
William J. Banks, 1981. Applied Veterinary Histology. Williams and Wilkins, Los Angeles, CA.
Hans Elias, et al. 1978. Histology and Human Microanatomy. John Wiley and Sons, New York, NY.
Don W. Fawcett. 1986. Bloom and Fawcett. A textbook of histology. W. B. Saunders Company,
Philadelphia, PA.
Don W. Fawcett. 1994. Bloom and Fawcett. A textbook of histology. Chapman and Hall, New York,
NY.
Arthur W. Ham and David H. Cormack. 1979. Histology. J. S. Lippincott Company, Philadelphia, PA.
Luis C. Junqueira, et al. 1983. Basic Histology. Lange Medical Publications, Los Altos, CA.
L. Carlos Junqueira, et al. 1995. Basic Histology. Appleton and Lange, Norwalk, CT.
L.L. Langley, et al. 1974. Dynamic Anatomy and Physiology. McGraw-Hill Book Company, New York, NY.
W.W. Tuttle and Byron A. Schottelius. 1969. Textbook of Physiology. The C. V. Mosby Company, St.
Louis, MO.
Leon Weiss. 1977. Histology Cell and Tissue Biology. Elsevier Biomedical, New York, NY.
Leon Weiss and Roy O. Greep. 1977. Histology. McGraw-Hill Book Company, New York, NY.
During the hour of this lecture the
average male produced 6.6 million
spermatozoa
 Spermatogenesis is Divided into
3 Main Events

Event Cell Type Duration

Spermatocytogenesis Spermatogonia 27 Days

Meiosis Spermatocytes 24 Days

Spermiogenesis Spermatids 23 Days

Combined Duration = 74 Days

 Spermatogenic Cycle
Cycle of seminiferous epithelium is the series of
changes in a given region of seminiferous
epithelium between two appearances of the
same developmental step.

Using spermiation as a reference developmental

step, the cycle would be all events that occur
between two consecutive spermiations.
 Similarities
College vs. Spermatogenesis

Duration of entire process is longer

than the cycle length

Multiple groups of participants develop

simultaneously

Attrition of participants reduce product

yield

Timing of entry of participants in

different groups create defined
stages of the cycle
Stages of the
spermatogenic
cycle
in humans
Human spermatogenesis: path followed through
spermatocytogenesis, meiosis, and spermiogenesis as
a given cell travels through five spermatogenic cycles
 Stage

A stage of the cycle of seminiferous epithelium is

defined by an association of spermatogonia,
spermatocytes, and spermatids whose
developmental age differs by a multiple of the
cycle length plus a common remainder.

The value of the remainder differs for each stage

of the cycle.
Stage (cycle of the seminiferous epithelium or spermatogenic cycle)
- man-made (man defined) divisions of the spermatogenic cycle
- cellular association of germ cells at defined developmental steps
- association of spermatogonia, spermatocytes and spermatids whose
developmental ages differ by a multiple of the cycle length plus a
common remainder (unique for a given stage)

Multiple Remainder Multiple Remainder

72/16 = 4 + 8 66/16 = 4 + 2
56/16 = 3 + 8 50/16 = 3 + 2
40/16 = 2 + 8 34/16 = 2 + 2
24/16 = 1 + 8 18/16 = 1 + 2
8/16 = 0 + 8 2/16 = 0 + 2

Stage II Stage VI
 Differences
College vs. Spermatogenesis
Multiplying component of
participants in
spermatogenesis

Continuous entry of
participants in
spermatogenesis
manifested by the wave of
spermatogenesis along
the tubular length
Wave of the seminiferous
epithelium (not in humans)

"The wave is in space what

the cycle is in time."

Horse
rat