Plant Water Relations: Absorption, Transport and Control Mechanisms

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Plant Water Relations: Absorption, Transport and Control

Mechanisms
Chapter · April 2012

DOI: 10.5772/33478 · Source: InTech
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5
Plant Water Relations: Absorption,

Transport and Control Mechanisms
Geraldo Chavarria1 and Henrique Pessoa dos Santos2
1The University of Passo Fundo
2Embrapa Grape & Wine
Brazil
1. Introduction
Although water is abundant on Earth - covering 71% of the total surface - its distribution is
not uniform and can easily cause restrictions in availability to vegetal production. At global
scale, these restrictions are easily observed in dry climates and can appear in other regions
which do not currently experience drought, as provided by the future backdrop of climate
change (IPCC, 2007).
The influences of water restriction on losses in the production and distribution of vegetation
on the terrestrial surface are significantly larger than all other losses combined which are
caused by biotic and abiotic factors (Boyer, 1985). This striking effect of water on plants
emerges from its physiological importance, being an essential factor for successful plant
growth, involving photosynthesis and several other biochemical processes such as the
synthesis of energetic composites and new tissue. Therefore, in order to characterise the
growth and productive behaviour of plant species it is essential to have an understanding of
plant water relations, as well as the consequences of an inadequate water supply. Broadly,
the water state of a plant is controlled by relative rates of loss and absorption, moreover it
depends on the ability to adjust and keep an adequate water status. This will be considered
throughout this chapter.
2. Absorption and water flow through plants

Independent of the species, plants require from the soil a water volume that overcomes its
metabolic necessities. Through the transpiration process plants transmit to the atmosphere
the majority of the water absorbed from soil (generally around 90%). From this perspective,
it is noted that the plant water requirements are defined primarily by the atmosphere
evapotranspirative demand, which is a predominately passive process. Figuratively, and
with some caveats, we can compare a plant water flow with the principles of oil flow in the
wick of an old fashion lampion (Fig. 1).
When it is fired, the oil that is burned on the upper extremity of the wick is quickly replaced by
new one that is situated just below, and so on - following the physical forces of interaction
between liquid and tissue - until reaching the level of the fuel reservoir, in the basal extremity
of the wick. Applying this example to the plant, the burning of oil can be analogous to the
106 Advances in Selected Plant Physiology Aspects
process of the loss of water vapour through their leaves, i.e. the transpiration, which is caused
by the pressure gradient of vapour between tissue saturated with water from the leaves and
air, the “dry” atmosphere. The variations in this pressure gradient of the vapour will define
the evaporative demand of the environment where the plant is. In the other extreme - where it
represents the liquid reservoir of the lampion - we have water content present in the soil. In
this scene, it is noted that the water flow through the plant is dependent on the energy formed
by the gradient of the water content that is established between the soil and the atmosphere.
However, we will see throughout this chapter that plants, unlike our lampion, can and must
modulate this gradient in order to survive the wide variations of water availability between
types of soils, weather and seasons.
Fig. 1. Schematic representation of water flow through the plant (arrows), by analogy with
the oil flow through the wick of an old fashion lampion.
2.1 Water potential

The water content in the soil, plants and atmosphere is usually described as water potential
(Ψw). This is based on the relation between the water content in the part of a system and
pure water at the same temperature and atmospheric pressure, measured in pressure units
(megapascal-MPa or bars-Bar). By definition, the potential of free pure water at atmospheric
pressure and at a temperature of 25°C corresponds to 0 (zero) MPa. The contrast in the
water potential between two points invariably determines the direction of water transport in
a system. More precisely, the water potential represents all the water pressure in a given
system and it is the sum of osmotic potential (Ψπ), matrix potential (Ψm), hydrostatic
pressure or the turgor potential (Ψρ) and the gravitational potential (Ψg).
The osmotic potential (Ψπ) is the chemical potential of water in a solution due to the
presence of dissolved substances (solutes). This is always negative because the water moves
Plant Water Relations: Absorption, Transport and Control Mechanisms 107
from one point with a lower concentration of solutes (for example, pure water) to a point
with a higher concentration. So, the higher concentration of the solutes at a point which
makes the system more negative will be the osmotic potential in this place. The water
potential can also be influenced by a charged surface - mainly by soil components and cell
walls - which compose the influence of the matrix potential (Ψm). In the soil, this influence
of the matrix is so great that water potential is assumed negligible and therefore equivalent
to the matrix potential. Concerning the potential of hydrostatic pressure (Ψρ), it is noted
that this component of the water potential can be positive or negative and it refers to the
physical pressure that water exerts on a given system. For example, if we observe a turgid
cell of a root cortex or a leaf mesophyll, the hydrostatic pressure is positive. However, in a
xylem vessel subjected to a stressful condition - in a transpiring plant - this component of
hydrostatic pressure is negative. Finally, we should emphasise that the gravitational
potential (Ψg) - ignored in most cases - is very important in studies of the water potential of
tree species, where plant height exerts a great influence on water flow. Considering that this
gravitational component fluctuates at a rate of 0.1 MPa for every 10 meters of vertical
displacement, it is suggested to consider if when plant height is 10 m or more.
2.2 Water dynamics in soil-plant-atmosphere system

From these components of water potential we return to our lampion scheme (Fig. 1) and
show how the potential can vary over the continuum soil-plant-atmosphere, exposing the
control points of each step of water flow from the soil to the atmosphere.
2.2.1 Soil water

The water potential in soil affects water reservoir and its availability for plants, hence it has
a large impact on plant growth and production. Furthermore, the soil water content exerts a
great influence on some physical and chemical properties of soil, such as the oxygen content,
which interferes with root breathing, microbial activity and soil chemical status. Water
potential is directly dependent on soil physical characteristics, and varies with time and
space, depending on soil water balance. That balance is determined by input (rain,
irrigation) and output of the soil (drainage, evaporation and root absorption). It is
noteworthy that the amount of rain affecting soil water reservoir is only the effective
precipitation. This is the amount of precipitation that is actually added and stored in the soil.
For example, during drier periods less than 5 mm of daily rainfall would not be considered
effective, as this amount of precipitation would likely evaporate from the surface before
soaking into the ground.
It is important to emphasise that behaviour of water into soil differs from that in a pot, like
the oil in the lampion reservoir (Fig. 1). That is, soil water interacts with the matrix and
solutes, and it is under pressure or tension, resulting in various energy states, relative to free
water (Kirkham, 2005). With regard to the physiological aspect, it is important to point out
that the water content in soil is associated with three terms: field capacity, the permanent
wilting point and the available water content.
The term “field capacity” corresponds to the maximum water content that a given soil can
retain by capillarity, after saturation and gravity drainage, and it is conventionally estimated
as the water content when the matrix potential is -0.03 MPa (-0.3 Bar). In spite of the great
applicability of this term to irrigation management, field capacity has been recognized as an
imprecise term due to theoretical advances and precise irrigation techniques. It is because
the capillary soil water constantly (even slowly) decreases (due to evaporation from soil
surface or drainage losses) and never stabilises (Fig. 2), it turn the soil water potential
decreases while the matrix potential increases. This is most evident with medium and fine
texture soils (for example, those rich in clay and organic matter), which maintain a
significant drainage rate over a long time. Therefore, there is no real and unique value for
accurately characterising the field capacity of a given soil. Furthermore, the continuous
drainage can induce an overestimation of the water consumption of the plant. Despite these
uncertainties, the term field capacity is still useful for a qualitative understanding - rather
than a quantitative understanding - of the water behaviour of a particular soil, providing an
estimate of the maximum limit of water accumulation. It is noteworthy that the inaccuracy
of the field capacity determination occurs mainly when analysis takes place on samples in
the laboratory, which can be contoured with evaluations directly in the soil, with specific
sensors and considering together all characteristics of each site. In general, clay soils or those
with higher content of organic matter (upper to 5% of organic matter) present a higher soil
water holding capacity (average field capacity ranging from 35 to 40%vol). In contrast,
sandy soils has a lower water holding capacity and field capacity typically ranges from 10-
15%vol. It is important to observe that field capacity cannot be regarded as a maximum limit
of the water available to plants, due the fact that plants also use free water that is in contact
with the roots at the moment of soil drainage.
The wilting point (WP) is another important parameter in soil water dynamics as it
dramatically affects plant physiology. This term is also known as the permanent wilting point,
and can be defined as the amount of water per unit weight (or volume) of soil that is so
tightly retained by the soil matrix that roots are unable to absorb causing the wilting of
plant. In others words, it corresponds to the water potential of soil under which plants
cannot maintain turgor pressure, even if a series of defence mechanisms have been triggered
(e.g. increased ABA synthesis, stomatal closure, osmotic adjustment, leaf fall) (For more
details see the Chapter by Mastrangelo et al.).
Similarly with FC, the value of water content in a soil at WP is not a unique and precise
value despite it is conventionally measure at -1.5 MPa (-15 Bar) (Fig. 2). The WP is
influenced by the physical and chemical characteristics of soil, but also by the plant species
considered. This is because various plant species differ in their ability to deal with low soil
water content due to differences in roots anatomy and depth, osmotic adjustment capacity
and other defence drought mechanisms.
Conventionally, the wilting point is estimated as the water content when the matrix potential of
the soil is -1.5 MPa (-15 bar). Nevertheless, some species of plants can absorb water soil at a
potential much smaller than this limit. For example olive trees can set a water potential
gradient between dry soil (-3 MPa) and leaf (-7 MPa) (Dichio et al., 2006). Similarly, Larrea
divaricata may absorb water at -6.0 MPa soil water potential (Kirkham, 2005). Another species
of the same genus of desert plant (Larrea tridentata) can survive with soil water potentials up to
-11.5 MPa, maintaining the photosynthetic activity of leaves within the range between -5 and
-8 MPa (Fitter & Hay, 2002). These examples serve to explain that the permanent wilting point
does not exclusively depend on the soil but also on the plant species. At the permanent wilting
point, the water potential of soil tends to be less than or equal to the osmotic potential of the
plant, which is extremely low in plants adapted to dry environments.
Fig. 2. Variation of the matrix of the water potential of soil (Ψsoil, negative values) in
relation to water volume (cm3.cm-3 of soil), characterising the limits of the field capacity (FC,
-0.03 MPa) and wilting point (WP, -1.5 MPa) of a given soil. The curve was generated from
results obtained by Santos, H.P., in Bento Gonçalves-RS, Brazil. 2010.
The indiscriminate use of a fixed value to estimate field capacity (FC) and the permanent
wilting point (WP) can generate false interpretations. However, this reference to the water
content in the soil is essential for calculating the available water content (AWC) for the plants.
The AWC is calculated considering the soil volume explored by roots and the % of water
content determined as the difference between FC and WP. Due to this interval of water
availability, one may assume that water could be absorbed by the roots with the same
facility in the range between FC and WP. For some plants this may be true, given that the
energy to extract water from the soil is small, compared to the energy needed to transport
the water from the root system to the atmosphere. However, with the reduction of soil water
potential, there is also a reduction in its hydraulic conductivity (i.e. water moves slowly in
the soil), limiting the water absorption capacity of the roots. In this scene - and for a majority
of crops - the yields are reduced if the water content in the soil approaches the wilting point.
Thus, the available water content should be considered as a relative value and, for the same
soil water potential, it may have different proportions of accessibility, depending on the
ability of each species to exploit or capture available water.
2.2.2 Water absorption by the roots

As was pointed out in Figure 1, the water flow of a plant is primarily controlled by the
transpiration rate. In this flow system it is essential indeed that there are no limitations on
water absorption by the root system. As the roots absorb water, there is a reduction in the
water potential in the soil that is in contact with the roots (rhizosphere). This process
establishes a water potential gradient between the rhizosphere and a neighbouring region of
the soil which presents a higher water potential and which coordinates the water movement
towards the roots of a transpiring plant (Fig. 3). This water movement in the soil occurs
mainly through mass flow due to the fact that the water filled micropores of the soil are
interconnected. Therefore, water flows from soil to root at a rate depending on the water
potential gradient between soil and plant which is affected by plant water need, hydraulic
conductivity of the soil, soil type and soil water content. Sandy soils have higher
conductivity due to greater porosity, but they also retain less water in relation to clay soils
or soils rich in organic matter.
At field capacity, water is initially removed from the centre of the largest pores (spaces ≥ 50 nm,
that are too large to have any significant capillary force) between the soil particles, maintaining
the water next to the particles due to adhesive forces. The reduction in water content causes a
drastic decrease in soil hydraulic conductivity, because the water is replaced by air in the
spaces between the soil particles (Fig. 3). Thus, the water movement in the soil is limited to the
periphery of soil pores, which can promote restrictions in the hydraulic conductivity to the
root surface and reach the permanent wilting point (discussed previously).
Soil particles
Symplastic and
Transmembrane
pathway
Endoderm
air
water in different
level of soil
interaction
Apoplastic
pathway
Fig. 3. Detail of the rhizosphere. Note the water adsorbed in soil particles. As the water is
absorbed by the roots, the open spaces filled with air increase (small arrows). During
absorption, water can flow by symplastic, apoplastic and transmembrane pathways
(detailed in the text) to the endoderm cells, where the Casparian strip is present
(represented by black points between the endoderm cells).
The water absorption by the roots is related to its surface directly in contact with soil. Thus,
longer and younger (less suberised) roots with more root hairs are essential for increasing
the contact surface and improve the water absorption capacity of the soil (Fig. 3). Moreover,
the distribution and proportion of the roots is very important for meeting the water demand
of a plant. In humid regions, as tropical rain forest, plants usually do not require very
extensive root systems (i.e. root:shoot ratio < 0.15, Abdala et al., 1998), because a small
volume of soil can meet the demands of transpiration. In addition, the water absorbed from
that small soil volume is frequently (and easily) replenished by rainfall. This condition in
turn induces a reduction of the root:shoot ratio. On the other hand, in dry regions, the
plants invest more in their roots, increasing the root:shoot ratio such that the roots can
represent upper to 90% of a plant biomass in some species of a desert climate, such as
observed in some species from open areas of the Bana woodland in southern Venezuela
(i.e. root:shoot > 5, Bongers et al., 1985) and from savanna in Brazil (Abdala et al., 1998). It
is important to note that the use of this root:shoot relation in the classification of plants
with respect to their habitat must be made with caution. In many species, a higher
investment in roots is more related to the accumulation of reserves and not specifically to
an increased root surface for water absorption (e.g. Manihot spp.). A higher investment in
roots can also support a process called hydraulic lift, when the roots translocate the water
from the soil positions with a greater water potential (for example, deeper) to soil
positions with a less negative water potential. This process promotes a hydraulic
redistribution (Burgess & Bleby, 2006) in the soil independently from plant transpiration,
because it occurs when the stomata are closed (e.g. at night in C3 and C4 plants, and
during the day in CAM - Crassulacean Acid Metabolism - plants).
With the water reaching the roots, the absorption process is directly dependent on the
water potential gradient between the rhizosphere and the root xylem. There are two ways
to establish this gradient, characterised by two absorption processes: 1) osmotically driven
absorption, common in plants with low transpiration activity; and 2) passive absorption,
which dominates in plants with high transpiration activity. The osmotically driven
absorption occurs in plants under conditions of heat and non-limiting water availability in
the soil, but with a restricted capacity of transpiration (for example, without leaves or
with a limited vapour pressure deficit). In these cases, there is an accumulation of solutes
in xylem vessels (for example, sucrose by degradation of starch reserves in the roots),
reducing the xylem water potential in relation to the soil water potential (Kramer &
Boyer, 1995). This condition results in water absorption and an increase in root pressure,
which is itself responsible for the guttation that means the leaf water output through the
hydathodes (pores located at the margins of the leaves) (Fig. 4). Moreover, increase in root
pressure also promotes the water exudation in lesions of branches, easily observed in
some species (e.g. exudation on branches of grape and kiwifruit plants after pruning in
early spring).
In passive absorption and with an increasing rate of transpiration, the tension in the xylem
vessels increases, indicating a predominance of the pressure potential influence upon the
osmotic potential through the establishment of the water potential gradient between the root
xylem and the rhizosphere. Under these conditions, the roots become a passive absorption
organ, where the water is sucked into a mass flow promoted by the transpiration activity of
aerial parts of the plant. A grapevine, for example, which during its annual growth and
production cycle transpires between 650 to 900 mm of water, in accordance to
environmental conditions where it is growing, and this volume corresponds to about 85% of
all its absorbed water (Mullins et al., 1992).
Fig. 4. Detail of a leaf of a wheat plant (Triticum aestivum L.) presenting guttation in the
morning. Photo: Ana Cláudia Pedersen.
The water intake in the roots can follow three ways into the root tissue in relation to the
route of the epidermis to the endoderm of the root, called radial water transport (Fig. 3): 1)
apoplastic, where the water moves through the intercellular spaces and does not pass
through any membranes, exclusively occupying the continuous network of the cell walls; 2)
symplastic, where the water moves exclusively from one cell to another through
plasmodesmata connections; and 3) transmembrane, which corresponds to a mixed path
between the first two, where the water goes in one direction through the root tissue,
entering (symplastic) and exiting (apoplastic) cells. The relative importance of these
pathways is still a cause of much discussion, but there is some evidence for the suggestion
that plants displaying low transpiration activity predominantly witness symplastic
transport, while those displaying high transpiration activity witness a greater proportion of
apoplastic transport (Boyer, 1985; Steudle, 2001). Another important detail in relation to
these different pathways is relevant only in the outer layers of the root tissue, because in the
endoderm the water apoplastic flow is limited due to the Casparian strip (Fig. 3). In this
hydrophobic barrier, the radial and transverse endodermal cell walls are impregnated with
lignin, suberin, structural wall proteins and wax. Note that in many plants this barrier also
occurs in the epidermal cells, forming a double layered hydrophobic barrier in the roots
(Enstone et al., 2003). It is important to note that the Casparian strip does not always
establish a barrier that is totally impermeable to water and solutes coming from the soil.
This can be observed in - for example - the development of young roots where pericycle
growth can break parts of the endoderm and allow free access to water until the
reconstitution of the tissue.
With regard to water absorption control in the roots, plants also present a family of
membrane water transporter proteins (water-channel proteins), called aquaporins. These
proteins have a critical role in water absorption, reducing the resistance to the water flow
along the transcellular path. The number of these proteins available for the root surface is
variable throughout the day, being higher during the photoperiod due to the higher
demands of photo-transpiration. The aquaporins are controlled by many endogenous and

exogenous factors of the roots, such as pathogens, phosphorylation, pH, solute gradient,
temperature and all environment factors that interfere in hydraulic conductance along the
water flow by the plant (Chaumont et al., 2005; Maurel et al., 2008).
2.2.3 Ascension of water through the plant: Vascular system

The presence of plants outside the water environment - among other factors - has been
related to the evolution of the vascular system, which allows for the speedy upward
movement of water to meet the demand of transpiration from the leaves. Water supply
through cells by diffusion (difference in chemical gradient) alone is not able to maintain the
hydration of a perspiring canopy plant. The need for a vascular system is more evident
when we observe the hydraulic dynamic of a tree during a hot day, which demands a large
flow of water (for example, 200 to 400 liters day-1) to fit a transpiring surface that is situated
along elevated positions, and in some species is higher than 100 meters (e.g. Sequoiadendron
gigantea).
The water flows from the roots to the shoot of the plant through the xylem. The general
mechanism to explain this upward movement of water is the cohesion-tension theory, which
was proposed in the late 19th century. Basically, this theory holds that the water evaporated
in leaves establishes a tensile strength in the xylem, where the hydrogen bonds provide a
continuous intermolecular attraction (cohesion) between the water molecules from the leaf
to the root (Fig. 1). Thus, the water column in the xylem lumen is driven out of a region with
a higher water potential, i.e. from the root and the stem, to a region with a lower water
potential, as the leaves, and finally toward to the air that can reach very low water potential
(e.g. -100 MPa, at 50% of air relative humidity).
Recently, the cohesion theory has been questioned as a result of assessments of tension in
xylem vessels, which do not present a direct relation with tension values measured on
leaves through pressure chambers. Furthermore, it is assumed that the hydrophobic
interaction between the internal walls of the xylem and the sap composition (lipids,
proteins, polysaccharides etc.) prevents the development of a tensile strength larger than 1
MPa (Zimmerman et al., 2004), which is smaller than the estimated tension of rising water in
a 30 m high tree (3 MPa). However, despite these questions, many studies argue that the
fundamentals of the cohesion-tension theory are still valid for explaining the water flow in the
continuous soil-plant-atmosphere (Richter, 2001; Steudle, 2001; Cochard, 2002; Tyree, 2003).
These elements support the idea that the water column of leaves to the roots provides an
auto-regulation mechanism between the process of loss and absorption of water by the
plants. Therefore, although the importance of the cohesion-tension theory has been neglected
by some critics, this mechanism is considered to be essential for the survival of plants
during the transpiration process, i.e. the loss of water.
With rising water in the trunk, in addition to pressure force, there is also capillarity
strength in the vessels. In a perspiring plant, the water moves continuously from the
xylem bundles to the intercellular spaces in the leaves, where the water potential is lower.
Due to capillarity strength, water which evaporates through leaf stomata is replaced by
the water contained in the lumen of the vascular bundles. In physiological temperatures
(25°C), the cohesive forces between the water molecules are sufficient to prevent the
disruption of the water column. This tension and the capillarity forces present in the
vascular bundles also present resistance to the water flow along the plant by two major
ways: 1) the inherent properties of the xylem flow and 2) the geometric aspects of the
xylem conduits (vessel elements and tracheids). In this respect, it is notable that plants with
the vessel elements of xylem can present a significantly lower hydraulic resistance than
plants with tracheids (Tyree & Zimmermann, 2002). As such, the xylem diameter has a
great influence on the hydraulic conductivity or water flow (Jv, mm s-1), according to the
Hagen-Poiseuille equation which describes the transport of fluids in ideal capillaries:
Jv = (πR4∆Ψ) / 8ηL (1)

In this equation, ∆Ψ is the difference in the water potential (MPa) between two points of
observation throughout the capillary, R is the radius of the capillary (mm) with a
determined length L (mm), through which occurs a flow with a constant viscosity η (1,002 x
10-3 Pa second-1, at 20°C) (Nobel, 2009). Accordingly, this equation shows that Jv is
proportional to the fourth power of the vessel conductor diameter (the xylem). According to
this logic, a trunk that presents few xylem vessels with large diameter has a greater Jv than a
trunk with the same xylem area distributed in a larger number of vessels of smaller
diameter. In general, xylem vessels with larger diameter are also longer. Xylem vessel
diameter is also variable throughout the growth season, being larger in those vessels formed
early in the growing cycle. This makes growth rings visible in transversal cuts of tree
species. The vessels diameter is an important factor in preventing cavitation or embolism
(formation of air bubbles by breaking the water column under high tensile strength, such as
values close to -30 MPa), with thinner vessels being less susceptible to such a water column
breakage. Generally, the cross-sectional area of vascular bundles is proportional to the
transpiring leaf surface. This may be observed in plants adapted to arid environments,
which have thinner vessels and a small transpiring surface, as reduced root:shoot ratio.
2.2.4 Leaf water and transpiration

Returning to our lampion scheme (Fig. 1), we emphasise that the leaves are the final frontier
of the water flow in the continuous soil-plant-atmosphere system. In leaf mesophyll there is
an extensive system of intercellular spaces - present in cell walls - which correspond to the
internal surface of water contact with the air. By this interface between the cell walls and the
intercellular spaces is established a water potential gradient, mobilising the water by the cell
walls from the final extremities of the xylem bundles. This water flow by the cell walls
occurs in an analogous way to the principles of water movement through the soil matrix,
and the water interacts with cellulose microfibrils and other hydrophilic components of the
cell wall (Fig. 5). Due to the high surface tension and as a result of water evaporation in the
surface of the cell walls which are in contact with the air in the intercellular spaces, it is
established that the tensile strength is transmitted to the xylem. Therefore, it is the tensile
strength that drives the upward flow of the water column from the root and is produced in
the internal evaporation process in the leaves. The maintenance of this process - as a result -
depends on the output (in the atmosphere) of water vapour present in the intercellular
spaces. As the leaf cuticle represents a barrier to water outlet - allowing on average only 5%
of water permeability - the water vapour moves from leaf intercellular spaces to the
atmosphere predominantly through stomatal diffusion. This process of the loss of water
vapour by the leaves is called transpiration and corresponds to the majority (90%) of the
volume of water absorbed by plants.
Transpiration has a number of positive effects (e.g. helps with mineral transport and leaf
cooling) however it also may contribute to induce water stress when soil dry.
In the continuum soil-plant-atmosphere of water flow, there are two major factors
determining the water potential of a plant: 1) the water potential of the soil, which
characterises the water supply; and 2) transpiration, which defines the loss of water. The
plant, which is an intermediate in this process, may regulate the water potential gradient
between the soil and the atmosphere primarily through the regulation of stomatal
conductance.
Water in cell wall
air layer limitrophe Water

vapor
Fig. 5. Leaf water. The evaporation from the cell walls of mesophyll should be noted. The
water vapour escapes through the stomatal opening and this flow is directly influenced by
the boundary layer of air.
The stomata have a quick and fine control of the water relations of a plant, coordinating the
control of the water potential gradient between the leaf and the air. In this interface of the
leaf with the environment, it is important to note that small changes in the relative humidity
of the air are reflected in major changes in the water potential gradient, which requires a
stomatal control so as to maintain the water stability of the plant. A simple variation from
100% to 99% of the relative humidity already corresponds to a decrease of -1.36 MPa in the
water potential of the air. This decrease becomes more evident in the water potential of the
air in average (80%) and in extreme (50%) conditions of relative humidity, which
respectively provide values of -14 and -93.6 MPa at 20°C (Nobel, 2009). If we compare the
water potentials of the air with the average water potential of a mesophyte plant (-0.5 MPa),
the high gradient always determines that the water is diffused from the leaves to the air
(Fig. 4). Throughout this water route between the leaf and the air there are two components
that can exert resistance to the diffusion process: 1) stomatal resistance, which is coordinated
by the stomatal opening; and 2) resistance of the air boundary layer, which is located closest to
the leaf surface (Fig. 5) and it is directly influenced by wind speed. The higher the speed of
the wind, the greater is the frequency of air renewal in this layer surrounding the leaf,
restricting diffusion resistance for the maintenance of a major gradient of the water
potential. Morphological and anatomical variations among leaves can interfere with the
speed of displacement of this thin layer of air, restricting the rate of transpiration in dry
environments. Among these modifications, we highlight the presence of hair, the stomata
located at the lower surface of the leaf, and the shape and size of the leaves. Although these
changes interfere directly with the transpiration rate, they do not exercise a variable and
instantaneous control in relation to the ambient conditions, as is done by the stomata.
During the day, great changes occur in the water potential along the soil-plant-atmosphere
system. Initially, let us consider a mesophyte plant in a constant atmospheric condition of
75% relative humidity at 20°C (-39 MPa of air water potential) and soil without water
restriction (at field capacity). During the nocturnal period, the transpiration is virtually nil, by
stomatal closure, promoting an equilibrium between soil, root and leaf water potentials
leading the potential gradient to be near zero (Fig. 6). With the first rays of the sun, at dawn,
the stomata opens, allowing the water diffusion of the leaf (transpiration) and, as a
consequence, reducing the leaf water potential.
Fig. 6. Schematic daily variation of soil (Ψsoil), root (Ψroot) and leaf (Ψleaf) water potential of a
plant well hydrated (A) and under water restriction (B).
When leaf water potential does not equilibrate with Ψsoil at the end of day (case B), the
permanent wilting point (Ψleaf wilting) is reached.
As a result - and with a delay that depends on the cohesion-tension forces of water and the
water column size between the leaf and root - it begins the reduction of the root water
potential. This reduction is slight (on average -0.3 MPa), due to high water availability of the
soil. The reduction of the water potential in the plant reaches the minimum limits during the
hottest times of the day, forcing the stomata to close for small intervals for extreme cases of
transpiration demand (Fig. 6A). At dusk, these variations of the water potential in the leaf
and the root are reversed.
In considering a plant under the same atmospheric conditions, but with severe water
restriction (a soil water potential near to the permanent wilting point), the water potentials
of the leaf and root necessarily reach values that are more negative for the sustainability of
the water flow by the plant. At high limits of negative potential, the differences among the
water potential of the leaf and root are smaller, but with great difference in relation to the
soil water potential (Fig. 6B). If this condition persists, there will be a decrease in the turgor
pressure of the leaves, causing temporary leaf wilting, which is recomposed during the
nocturnal period. Mesophytic plants can tolerate this reduction in soil moisture up to the
limit of -1.5 MPa, while some xerophyte plants can reach limits of -5.5 MPa (Nobel, 2009).
3. Physiological and biochemical aspects of water in plants

In a plant cell, the water is predominantly located in the vacuole and represents the majority
of the mass of growing tissue (on average 90%). This predominance is due to the importance
that this universal solvent represents in the physiological and biochemical processes of a
plant cell. At the cellular level - due to its polar structure - the water acts on the dissolution
and mobilisation of ions and organic metabolites, such as amino acids, proteins,
carbohydrates and hormones. These water properties are also crucial for the flow between
parts of the plant, acting directly in the transportation of nutrients, carbohydrates and
hormones. The water acts in membrane integrity and the support of herbaceous plants
through the cells turgor pressure, which varies between 1 and 5 MPa. This turgor pressure is
also essential in the expansion process of cell walls and in growing tissues. This subsection
will expose some physiological and biochemical aspects of the water relations in plants.
3.1 Water deficit and its effects on plant growth

When a plant is under water deficit, responses at physiological, biochemical and molecular
scale are triggered (Shao et al., 2008). Physiological responses are linked to a condition of
recognition of stress by the root system, turgor changes and water potential and,
consequently stomatal conductance, internal CO2 concentration and photosynthetic activity
decrease. In biochemical terms, there will be a decrease in the photochemical activity of
photosynthesis, rubisco enzyme activity and the accumulation of secondary metabolites
linked to stress (such as glutathione and polyamines). From a molecular perspective, several
genes expressed under stress conditions are activated, such as genes linked to the
biosynthesis of abscisic acid and the synthesis of specific proteins.
As the cells hydration is reduced and the plant goes into a condition of water deficit, abscisic
acid and solutes increase in the plant, especially in the root system (the increase in solutes
occurs in a relative manner, due to water reduction). These factors will reduce stomatal
conductance and, consequently, photosynthetic activity which ultimately will result in a
reduction in the synthesis of proteins and cell walls, as well as a decrease in the rate of cell
expansion (Taiz & Zeiger, 2004). The sum of these responses to water deficit contribute to
explain the reduction of plant growth.
3.2 Elements that define water demand

Basically, according the cohesion-tension theory already described, the water demand of
plants is generated by a gradient at the top of the plant which generates negative hydrostatic
pressure that “suck” soil water through the plant into the atmosphere. Two main
environmental factors will determine this evaporative demand: wind speed and solar radiation
(Chavarria et al., 2009). Thus, those plants that exist in environments with high winds (e.g.
20 km h-1 or more) or intense solar radiation (e.g. 2500 µE m-2 s-1 of photosynthetically active
radiation) will suffer a greater water loss to the atmosphere. These plants need to make use
of water control mechanisms in order to tolerate these environmental conditions. Plants can
suffer morphological and anatomical alterations and osmotic regulation to improve stomatal
resistance and increase water absorption through the root system.
3.3 Mechanisms of water status regulation

3.3.1 Morphological and anatomical characteristics associated with water control
Plants live with a constant dilemma, namely to undergo photosynthesis while preventing
water loss. Every time plants open their stomata to allow the influx of carbon dioxide, they
lose water by diffusion.
When plants achieved their “terrestrial status” and left behind the conditions of algae, they
required several changes to adapt to this new environment. They acquired a root system for
physical support and absorption of water and nutrients, vascularity for the movement of
water and photoassimilates, and a stomatal complex and a wax layer for the regulation of
water loss to the atmosphere.
It is known that intense solar radiation is accompanied by elevated temperatures, which can
cause severe physical damage to leaves and lead to senescence and leaf abscission or, on
small scale, reduce carbon assimilation. This situation forces plants to adopt strategies to
minimise such negative effects.
In order to adjust the balance between water availability and atmospheric demand, plants
can reduce the size and number of leaves. The organisation of the mesophyll and leaf
dimensions is modified by water restrictions, and can provide a strategy for plants to affect
the stomatal conductance and CO2 diffusion (Evans et al., 2009). Plants which need greater
efficiency in water use (μmol CO2 fixed / H2O transpired) have a strategic demand to retain
higher CO2 concentrations inside. Characteristics of leaf anatomy related to water deficit are
reduction of thickness, higher cell density and smaller intercellular spaces, all of which try
to mitigate the problems of excessive water loss (Chartzoulakis et al., 2002). The reduction in
cell size and, consequently, the reduction of tissues are associated with the turgor decrease
of cells (Ogbonnaya et al., 1998). However, smaller cells may stay more turgid when
compared to larger cells, having better capacity to tolerate conditions of water restriction
(Burghardt et al., 2008). These changes in cell size result in an increased internal surface to of
CO2 exchange per leaf area, seeking to maintain the photosynthetic rate with reduced
stomatal conductance (Ennajeh et al., 2010; Syvertsen et al., 1995).
The relation of the root with shoots is also strongly modified due to conditions of water
unavailability. One way that plants use to keep their water status stable is by reducing the
growth of the shoot, in order to reduce leaf area and water loss to the atmosphere. At the
same time, plants can invest in the growth of the root system, in order to increase the soil
volume explored and in turn water absorption. All these mechanisms occur through
hormonal changes as initiated by abscisic acid signals. Nevertheless, later on it will be the
effects of interactions between auxins, cytokines and gibberellins which define the relation
between root and shoot.
Anatomically, the change in vascular diameter may be a response to water deficit conditions
and tends to decrease under water deficit (Kutlu et al., 2009). By reducing the radius of
vessels, xylem conductivity is reduced (increased resistance) according to equation 1. There
are some situations of high evaporative demand which increase the tension in the xylem
vessels, causing a disruption of the water column and the formation of air bubbles by
embolism (Tyree & Sperry, 1989). The disruption of the water column can also occur under
conditions where the water freezes inside the plant.
Another mechanism which protects the leaf from attack by insects and helps the plant to
avoid water loss is the presence of leaf trichomes (Molina-Montenegro et al., 2006).
Trichomes reduce water loss by: 1) reducing the arrival of solar radiation on the leaf surface;
and 2) reducing the interference of wind on the boundary layer, which reduces the
differences in water potential between the leaf and the atmosphere (Ehleringer, 1984;
Vogelmann, 1993).
Leaf cuticle presents variations in anatomical position and chemical composition.
Chemically, these cuticles are characterised by two specific groups of lipids: 1) cutin, which
forms the support structure of the membrane; and 2) waxes deposited on the external
surface - called epicuticular waxes - and also strongly dispersed within the matrix of the cutin
(below the surface) called cuticular waxes (Devine et al., 1993). The most important function
of epicuticular waxes is to avoid leaf water loss to the environment. In addition, they reduce
leaf nutrient loss, prevent excessive solar radiation, pathogenic microorganisms, cooling,
wind damage and physical abrasion (Vigh et al., 1981; Mendgen, 1996; Barnes & Cardoso-
Vilhena, 1996; Scherbatskoy & Tyree, 1990; Kerstiens, 1996; Eglinton & Hamilton, 1967).
Epicuticular waxes may be amorphous, have a flat format covering the entire surface area of
the leaf, or have the shape of a crystal or blade. Involved in the formation of these waxes are
alkanes, esters, ketones and alcohols (Shepherd et al., 1995).
An important aspect of plant morphology is the density of leaf veins. Angiosperms average
8 mm of vein per mm² of leaf area, while non-angiosperms have consistently averaged close
to 2 mm mm-2 throughout 380 million years of evolution (Boyce et al., 2009). This was an
important ecological strategy for the increment of photosynthesis, especially under
conditions of higher temperatures and transpiration rates.
The most important anatomical tools to optimise the plant water use is the stomatal
complex. Variations in opening, size and position of the stomata in the leaf help plants grow
under conditions of water deficit (Larcher, 1995).
3.3.2 Stomata metabolism

Stomatal complexes are present in green stems, flowers and fruits, averaging 30 to 400 per
square millimetre. They are formed by an opening, called a pore; two guard cells that are
responsible for opening and closing the pore; and in some situations neighbouring cells
called subsidiary cells, whose function is to support the guard cells. The stomata liberate into
the atmosphere around 95% of the water that exits from the leaves and causes 90% of gas
exchange (in this case, also involving carbon dioxide), while only 5% of the water output is
through the cuticle. There are two types of stomata morphology: kidney-shaped and grass-
like, sized between 5 and 15 μm wide and 20 μm long (Fig. 7).
Guard cell turgid

Guard cell turgid
Subsidiary cell
Guard cell flacid
Pore
Pore
Guard cell flacid

Epidermal cell
Epidermal cell
A B
Fig. 7. Stomatal complexes: grass-like (A) and kidney-shaped (B), with the left guard cells
turgid (open pore) and the right flaccid (closed pore).
The stomata works as a hydraulic valve regulated by water. When turgid, the stomatal pore
opens, and when flaccid due to water loss it closes (Fig. 7). In the early morning, light in the
blue band (440-490nm) indicates the arrival of solar radiation and possibility of
photosynthetic activity. Thus, the plant opens the stomatal pores to allow influx of carbon
dioxide. The first theory about stomatal opening control was called the “starch-sugar
hypothesis,” and it was widely accepted during the early 1940s. This hypothesis suggested
that the hydrolysis of starch in soluble sugars decreased the osmotic potential of cell,
promoting water absorption and stomatal opening. However, that theory lost strength as
advances were made by studies of potassium movement in the guard cells. Currently, three
modes covering the osmoregulation processes of guard cells are accepted: 1) the influx of
potassium and chloride through proton pump activation and the synthesis of malate from
starch breakdown; 2) sucrose synthesis by starch hydrolysis; and 3) sucrose synthesis by
carbon dioxide fixation due to photosynthetic activity.
The blue light signalling process occurs through receptor pigments (phototropins).
Thereafter, the opening process occurs by the reduction in the osmotic potential of guard
cells by potassium input, consuming ATP by ATPase located in the plasmatic membrane.
This ATPase release protons inside the guard cells, causing a variation in pH around 0.5 to
1, which enables membrane hyperpolarisation and, consequently, the opening of channels
favours potassium absorption. The resulting osmotic gradient causes water movement
towards these cells. Therefore, the guard cells become turgid and the stomatal pore opens,
due to the action of cellulose microfibrils (Fig. 7). However, it is important to emphasise that
chloride and malate also contribute to the establishment of the osmotic gradient in guard
cells exposed to blue light.
When solar radiation begins to decrease this affects the photochemical phase reducing levels
of ATP and NADPH+ resulting in losses for the biochemical phase. Internally, the carbon
dioxide content begins to rise as they are not being used in the carboxylation process in the
Calvin-Benson cycle (C3). At the same time, calcium ions play an important role when they
enter the guard cells, causing solute output and decreasing the osmotic potential of these
cells, making the stomata close.
An atmosphere enriched with carbon dioxide can favour the photosynthetic activity of
plants with the C3 mechanism. However, excessive carbon availability will result in stomatal
closure in some species, even with a C3 mechanism (Ainsworth & Rogers, 2007). An example
of natural CO2 enriched conditions, where the plants can be submitted to excessive
concentrations of carbon dioxide, occurs near volcanic activity (Miglietta & Raschi, 1993).
3.3 Hormonal and molecular responses in different water conditions

The plants that display a higher production capacity, due to the morphological,
physiological and metabolic changes of their organs and cells, tend to present higher
demands on available resources and, consequently, possess greater vulnerability to
conditions of water restriction. However, there are several strategies for the adaptation to
dry environments that can be considered as a tool for progress in overcoming limiting
conditions on growth and production. Generally, some plants can accumulate water to
delay or escape such stress conditions, while others can deal with the stress through
decreased metabolic activity (Bartels, 2005). The effect of water restriction depends on the
degree and duration of the stress, the stage of plant development, the genotypic ability of
the species and environmental interactions. In recent years, and mainly due to climate
changes, several studies have looked to understand the biochemical and molecular basis
of water stress (Yokota et al., 2006; Xoconostle-Cázares et al., 2010). Water stress can
influence a plant on several levels, with cell expansion and growth being the first
processes to respond to water limitation. With the gradual increase of stress, other
processes are also affected, such as photosynthesis and allocation of assimilates. At the
cellular level, membranes and proteins can be damaged with the increment of reactive
oxygen species or peroxidation. These responses are common with other abiotic stresses,
such as the effects of salinity and low temperatures, resulting in the synthesis of a similar
group of proteins (Artlip & Wisniewski, 2002).
Tolerance to water deficit can be manifested in four ways: 1) the seasonal adjustment of
growth to avoid stressful conditions; 2) morphological adaptations, such as an increase in
the root:shoot ratio, a reduction in the leaf area and wax accumulation on the leaf surface; 3)
physiological adaptations, such as stomatal responses and leaf abscission; and 4) metabolic
changes. Among the metabolic changes of adaptation, osmotic adjustment represents the
most common change and results from the accumulation of certain metabolites.
The root has been generally accepted as the organ which acts on the perception of water
stress, although so far there is no knowledge about how the cells in roots perceive the soil
moisture content. However, when the plant water potential is influenced by the water
restriction of soil or saline stress, the stomata must respond quickly to avoid water deficit.
Signalling between the root and stomata is carried out by abscisic acid (ABA), which has
been considered to be a “plant hormone of stress” due to its participation in the signalling
networks of other factors of stress (Artlip & Wisniewski, 2002). ABA is synthesized from
carotenoid by the synthesising enzyme of ABA (zeaxanthin epoxidase, 9-cis-
epoxycarotenoid dioxygenase and aldehyde oxidase) which is induced in the root apex or in
the parenchyma cells of vascular bundles by water or saline stress. After the synthesis of
ABA in the roots, it is transported through the xylem to the leaves alone or conjugated with
glucose (the latter being more appropriate for transport over long distances). The proportion
of each form of transport (alone or conjugated) is variable between species (Sauter et al.,
2002). Once in the leaf, conjugated ABA is hydrolysed into its free form by the apoplastic
enzyme β-D-glucosidase, inducing stomatal closure through a signalling system in the
guard cells of chloroplasts (Yokota et al., 2006).
In addition to this long distance signalling between the roots and the stomata, recent
studies also point out that the leaves must act as sensors of relative humidity in order to
avoid desiccation. This is clear from observations of the extremely quick closure of
stomata with increments in the vapour pressure gradient between the leaf and the air,
even when there is adequate water availability in the soil (Assmann et al., 2000). Recent
works expose the possibility that leaf sensors of relative humidity are located in or near
their own stomata guard cells (Yokota et al., 2006). In addition, there is genetic evidence
that leaf sensitivity to relative humidity (RH) is related to ABA metabolism, exerting a
hormonal effect over a short distance. A recent study on genetic selection, based on
infrared thermal imaging, identified two genes (OST1 and ABA2) that are directly
involved in the signalling route of RH sensing in guard cells (Xie et al., 2006). OST1 codes
a protein kinase that is involved with stomata closure, while ABA2 codes an enzyme
involved in ABA biosynthesis. This reinforces the involvement of ABA as a mediator in a
signalling network of guard cells, which can be shared between different stimuli to
control stomata closure (Yokota et al., 2006).
ABA has been related to quantitative and qualitative variations in the gene expression and
protein synthesis stimulated by water stress (Artlip & Wisniewski, 2002). Meanwhile, it is
notable that some of the proteins that are de novo synthesized do not appear in the responses
to the application of abscisic acid (ABA) and these signals/response routes to the water
stress have ABA-dependent and ABA-independent routes (Yokota et al., 2006). In ABA-
independent routes, the signal molecule provided by the roots is still unknown.
Bioinformatics analysis has promoted advances in the identification of several factors of
transcription that are induced by water deficit, classified in six major groups: AP2/ERF
(APETALA2/ethylene-response factor); bZIP (Basic leucine-zipper protein); MYB/MYC
Zinc-finger protein; CDT-1; NAC and Dreb (Xoconostle-Cázares et al., 2010). In ABA-
dependent routes, the promoters of genes containing a cis-sequence of six nucleotides are
known as the ABA response element (ABRE). The genes’ expression of ABA-dependence is
activated by the AREB/ABF link - a transcription factor of the type bZIP - on the ABRE
sequence. Further, in a gene ABA-dependent RD22 expression there is involvement with the
transcription factors MYB and MYC, and they are related to the final stages of responses to
drought that are ABA dependent (Yokota et al., 2006). The promoters of genes related with
responses to drought also have an alternative regulatory sequence of nine nucleotides called
a dehydration response element (DRE). The DREs are involved in the ABA-independent
expressions of genes that are induced under drought conditions. The trans-factors to the cis-
elements are CBF/DREB1 and DREB2, which are expressed transiently after the detection of
drought and thereby stimulate the targeted genes involved in drought tolerance (Taiz &
Zeiger, 2002; Yokoda et al., 2006).
Genes that are stimulated by drought can be categorised into two groups: 1) coding genes
of those proteins responsible for protecting cells and organs against stress; and 2) coding
genes of those proteins necessary for signals’ translation and regulation of gene
expression. The proteins of the first group act directly on membrane functions, the
maintenance of water potential, proteins’ protection and oxidative stress control. Stands
out in this group the family of the Embryogenesis Abundant protein (LEA), which is formed
by five types of proteins based on the structural domain and which are suspected of
acting to protect the cell membrane (Taiz & Zeiger, 2002). Beyond that, and by their
hydrophilic properties, they act in water retention and prevent the crystallisation of other
proteins and molecules during drying. Within the LEA family itself, the D-
11/RAB/Dehydrins group stands out, whose function has been related with the
stabilisation of proteins and membranes. This group presents a wide distribution between
plant species, and can be considered to be an alternative to the constitutive defence
against rapid changes in the water status of tissue (Artlip & Wisniewski, 2002). The
aquaporins family represents another important protein in cell protection against water
stress, facilitating water absorption by the plasmatic membrane. Its importance in water
relations was recently evidenced by the differential accumulation of aquaporins in
relation to the degree of drought tolerance in varieties of beans (Montalvo-Hernández et
al., 2008). With regard to membrane protection and the water status of the plant we may
also highlight lipid transfer proteins (LTPs) that catalyse the transfer of several classes of
phospholipid and glycoproteins for deposits in cell walls or between membrane vesicles
(Kader, 1997). The results with LTPs show that these proteins are induced during different
conditions of stress and that they can act to increase membrane fluidity, decreasing water
loss by increasing tissue impermeability, and as a physical barrier to biotic stress (Treviño
& O´Connell, 1998; Maghuly et al., 2009; Gong et al., 2010).
Another protein family has been associated with responses to drought, namely Heat
Shock Proteins (HSPs), which are widely distributed in nature. These proteins are known
as molecular chaperones, acting in the folding and assembly of functional proteins and in
the removal of non-functional proteins, facilitating the recovery of cellular functions after
stress. Several HSPs - classified according to their molecular weight - are induced in
conditions of water and saline stress, such as HSP70 (the DnaK family), the chaperones
GroEL and HSP60, HSP90 and HSP100 and the small HSP (sHSP) (Alamillo et al., 1995;
Campalans et al., 2001, Wang et al., 2004). Within these proteins, there is the cyclophilin,
which is a chaperone protein with systemic properties and which is highly induced
during water stress, conferring multiple tolerances to abiotic stress (Gottschalk et al., 2008;
Sekhar et al., 2010). During conditions of stress, the recycling of macromolecules which
lose their function to maintain cellular homeostasis is essential. In this process, and under
conditions of water stress, an increase of protease activity has been observed (Campalans
et al., 1999; Seki et al., 2001), which is important in the destruction of denatured proteins
and in the recycling of amino acids necessary to synthesize proteins for water deficit
responses. In addition, the ubiquitin and polyubiquitin proteins are also induced with
water restriction, both of which act marking proteins for proteolytic degradation
(Campalans et al., 1999, Barrera-Figueroa et al., 2007).
3.4 Osmotic regulation

As discussed above, the osmotic regulation process of stomatal opening occurs by the
movement of solutes - called osmolytes - thereby influencing water movement between the
cells. This water movement has several purposes in relation to cell hydration inside plants,
such as the stomatal opening and higher water absorption via root system. Several
compounds that work in the osmotic regulation of plants are known: carbohydrates
(sucrose, sorbitol, mannitol, glycerol, arabinitol, pinitol) (Hare et al., 1998), nitrogen
compounds (proteins, betaine, glutamate, aspartate, glycine, choline, putrescine, 4-gamma
aminobutiric acid) (Kinnersley & Turano, 2000) and organic acids (malate and oxalate) and
so on (Sairam & Tyagi, 2004).
From the standpoint of thermodynamics, there are concepts relating to water potential that
are elementary in the explication of water movement. In turgid cells the water potential
(Ψw) is composed by pressure (Ψρ ) and osmotic (Ψπ) potentials. The greatest possibility of
interference in the Ψw of plant cells is by varying the amount of internal solutes. Thus,
osmotic regulation is a process in which variation of the amount of solutes - and
consequently in osmotic potential - interferes in water movement.
The higher the concentration of solutes in a solution, the greater will be the disorder of the
system (entropy). Furthermore, this condition leads to more negative osmotic potential and,
consequently, to a more negative water potential. The water inside the plant flows from the
sites with higher water potentials to those with lower ones. Thus, an increase in solute
concentration in the cell favours the entry of water.
The solutes that plants use to decrease the water potential and control the water flow must
have a low molecular weight. Among these solutes, the best-studied is potassium (K+), the
lack of which is related to tissue dehydration. However, most references point out that the
main role of potassium within the osmoregulation process is the regulation of stomata
opening (Läuchli, 1984; Hsiao & Läuchli, 1986). It also acts in water absorption by the roots,
in transpiratory control and in the cells’ ability to resist low temperatures (Grewal & Singh,
1980). In particular, under conditions of potassium deficit, sugars from starch hydrolysis
become more relevant in osmotic regulation than in the process of stomatal opening
(Poffenroth et al., 1992).
The proline amino acid also stands out as an osmotic regulator, and is linked to stress both
by water deficit and salinity (Molinari et al., 2004; Zhu et al., 2005). Nevertheless, proline
does not only exert the function of osmoregulation in plant cells during periods of water
deficit, but it can also protect against the activity of free radicals, regulate the pH in the
cytoplasm, protect against the denaturation of macromolecules and also act as a source of
carbon and nitrogen under conditions of stress (Sharma and Dietz, 2006; Vanrensburg et al.,
1993; Sivakumar et al., 2000; Díaz et al., 1999). Another compound with importance in
osmotic regulation is glycinebetaine, which, like proline, does not just exert or exercise its
function in osmotic potential. These compounds also act in the reduction of reactive oxygen
species (ROS) produced under conditions of stress in plants. With the process of tissue
dehydration, there is production of ROS, such as the singlet oxygen (1O2), the superoxide
radical (O2-), hydrogen peroxide (H2O2) and the hydroxyl radical (OH-) (Liu & Huang,
2000). Reactive oxygen species (ROSs) are produced from oxygen metabolism and play an
important role as indicators in the stress process of water deficit. Among these are oxygen
ions, peroxides and free radicals, and these compounds will cause oxidative stress in cells
and prejudice their operation.
4. Practical aspects of measuring plant water status

4.1 Pression chamber
The most widely used tool for determining water potential is the pressure chamber
(Scholander et al., 1965). The measurement is made on the section between a stem or a
branch and a vegetative part, which is placed inside a pressure chamber filled with an inert
gas (Fig. 8).
Fig. 8. Pressure chamber used to determine water potential in plants. Photo: Geraldo
Chavarria.
After leaf preparation, leaf bagging and chamber sealing, the chamber pressure is increased
gradually, resulting in a force inverse to the entry of water, in order to expel water through
the observed section of the xylem. Generally, the evaluation of the potential requires the use
of a magnifying glass. The pressure exercised to expel water is considered the pressure
which is retained in the cells. This technique is used in research for the characterisation of
the water status, but it may be useful in some crops as a tool for determining the appropriate
time for irrigation. Furthermore, some works also point to a high correlation between leaf
temperature measured with an infrared thermometer and water potential in the leaf
measured with a pressure chamber (Lafitte & Courtois, 2002).
4.2 Porometry
Another technique widely used to measure the water availability in plants is that of stomatal
diffusive resistance, using a device called a porometer, since the main route of gas exchange
between the plant and atmosphere occurs through the stomata. The resistance to water
diffusion by the stomata is measured in both sides of the leaf. The equipment may evaluate
four different processes: 1) mass flow (air forced through the leaf); 2) vapour diffusion (dry
air pumped into the chamber, which is equipped with a sensitive device to detect variations
in humidity inside the chamber); 3) maintenance flow (dry air pumped into the chamber
and steadily retained, where the flow changes are used to calculate resistance to stomatal
diffusion); and 4) state of equilibrium (to monitor the time necessary for the occurrence of
equilibrium by applying a dry air flow in the chamber attached to the leaf) (Fig. 9).
Fig. 9. Stomatal conductance determination using a porometer. Photo: Ana Cláudia Pedersen.
5. Acknowledgment
The authors would like to acknowledge the images of Figures 1, 3 and 4, which were
provided by Luciana Mendonça Prado, and 7 and 8 which were provided by Claudinei
Crespi. Also thanks to Ana Cláudia Pedersen for the support in the translation and for
providing Figures 4 and 9 and to Flavio Bello Fialho for the English review.
6. References
Abdala, G.C.; Caldas, L.S.; Haridasan, M.; Eiten, G. (1998). Above and belowground organic
matter and root:shoot ratio in cerrado in central Brazil. Brazilian Journal of Ecology,
Vol. 2, No. 2, pp. 11-23.
Ainsworth, E. A. & Rogers, A. (2007). The response of photosynthesis and stomatal

conductance to rising (CO2): mechanisms and environmental interactions. Plant,
Cell and Environment, Vol. 30, Nº 1, pp. 258-270.
Alamillo, J.; Almogura, C.; Bartels, D. & Jordano, J. (1995). Constitutive expression of small
heat shock proteins in vegetative tissues of the resurrection plant Craterostigma
plantagineum. Plant and Molecular Biology, Vol.29, pp. 1093-1099, ISSN 0167-4412
Artlip, T.S. & Wisniewski, M. E. (2002). Induction of Proteins in Response to Biotic and
Abiotic Stresses, In: Handbook of Plant and Crop Physiology. M. Pessarakli, (Ed.), 657-
679, Marcel Dekker Inc., ISBN 0-8247-0546-7, New York.
Assmann, S.M.; Snyder, J.A. & Lee, Y.R.J. (2000). ABA-deficient (aba1) and ABA-insensitive
(abi1-1, abi2-1) mutants of Arabidopsis have a wild-type stomatal response to
humidity. Plant Cell and Environment, Vol.23, No. 4, pp. 387-395, ISSN 1365-3040
Barnes, J.D. & Cardoso-Vilhena, J. (1996). Interactions between electromagnetic radiation
and cuticle. In: Plant cuticles: an integrated functional approach. G. Kerstiens, (Ed.),
157-174, Bios Scientific Publishers, ISBN 1-85996-130-4, Lancaster
Barrera-Figueroa, B.; Pena-Castro, J.; Acosta-Gallegos, J.A.; Ruiz-Medrano, R. & Xoconostle-
Cazares, B. (2007). Isolation of dehydration-responsive genes in a drought tolerant
common bean cultivar and expression of a group 3 late embryogenesis abundant
mRNA in tolerant and susceptible bean cultivars. Functional Plant Biology, Vol. 34,
No. 4, pp. 368-381, ISSN 1445-4408
Bartels, D. & Sunkar, R. (2005). Drought and salt tolerance in plants. Critical Reviews in Plant
Sciences, Vol.24, No. 1, pp. 23-58, ISSN 1549-7836
Bartels, D. (2005). Desiccation tolerance studied in the resurrection plant Craterostigma
plantagineum. Integrative and Comparative Biology, Vol. 45, No. 5, pp. 696-701, ISSN
1540-7063
Bongers, F.; Engelen, D.; Klinge, H. (1985). Phytomass structure of natural plant
communities on spodosols in southern Venezuela: the Bana woodland, Vegetatio,
Vol.63, pp. 13-34.
Boyce, C.K.; Brodribb, T.J.; Feild, T.S. & Zwieniecki, M.A. (2009). Angiosperm leaf vein
evolution was physiologically and environmentally transformative. Proceedings of
the Royal Society B, Vol. 276, pp. 1771–1776, ISSN 1471-2954
Boyer, J.S. (1985). Water transport. Annual Review of Plant Physiology, Vol.36, pp. 473-516,
ISSN 0066-4294
Burgess, S. & Bleby, T. (2006). Redistribution of soil water by lateral roots mediated by stem
tissues. Journal of Experimental Botany, Vol. 57, No. 12, pp. 3283-3291, ISSN 002-0957
Burghardt, M.; Burghardt, A.; Gall, J.; Rosenberger, C. & Riederer, M. (2008).
Ecophysiological adaptations of water relations of Teucrium chamaedrys L. to the
hot and dry climate of xeric limestone sites in Franconia (Southern Germany). Flora.
Vol. 203, pp. 3–13, ISSN 0367-2530
Campalans, A.; Messeguer, R.; Goday, A. & Pagès, M. (1999). Plant responses to drought,
from ABA signal transduction events to the action of the induced proteins. Plant
Physiology and Biochemistry, Vol. 37, No. 5, pp. 327–340, ISSN 0981-9428
Campalans, A.; Pagès, M. & Messeguer, R. (2001). Identification of differentially expressed
genes by the cDNA-AFLP technique during dehydration of almond (Prunus
amygdalus). Tree Physiology, Vol. 21, No. 10, pp. 633-643, ISSN 0829-318X
Chartzoulakis, K.; Patakasb, A.; Kofidisc, G.; Bosabalidisc, A. & Nastoub, A. (2002). Water
stress affects leaf anatomy, gas exchange, water relations and growth of two
avocado cultivars. Scientia Horticulturae, Vol. 95, pp. 39–50, ISSN 0304-4238
Chaumont, F.; Moshelion, M. & Daniels, M.J. (2005). Regulation of plant aquaporin activity.
Biology of the Cell, Vol. 97, No. 10, pp. 749-764, ISSN 0248-4900
Chavarria, G.; Cardoso, L.S.; Bergamaschi, H.; Santos, H.P. das; Mandelli, F. & Marodin,
G.A.B. (2009). Microclimate of vineyards under protected cultivation. Ciência Rural,
Vol. 39, No. 7, pp. 2029-2034, ISSN 0103-8478
Cochard, H. (2002). A technique for measuring xylem hydraulic conductance under high
negative pressures. Plant, Cell and Environment, Vol. 25, No. 6, pp. 815–819, ISSN
1365-3040
Devine, M.; Duke, O. S. & Fedtke, C. (1993). Physiology of Herbicide Action. Englewood Cliffs,
NJ: P.T.R Prentice–Hall.
Díaz, P.; Borsani, O. & Monza, J. (1999). Acumulación de prolina en plantas en respuesta al
estrés osmotico. Agrociencia, Vol. 3, Nº 1, pp. 1-10.
Dichio, B.; Xiloyannis, C.; Sofo A. Montanaro G. (2006). Osmotic regulation in leaves and
roots of olive tree (Olea europaea L.) during water deficit and rewatering. Tree
Physiology, 26:179–185.
Eglinton, G. & Hamilton, R.J. (1967). Leaf epicuticular waxes. Science, Vol. 156, No. 3780, p.
1322-1335, ISSN 0036-8075
Ehleringer, J.R. (1984). Ecology and ecophysiology of leaf pubescence in North American
desert plants. In: Biology and Chemistry of Plant Trichomes, E. Rodriguez, P.L. Healy
& I. Mehta, (Eds.), 113-132, Plenum Press, ISBN 030-6413-93-0, New York.
Ennajeh, M.; Vadel, A.M, Cochard, H. & Khemir, H. (2010). Comparative impacts of water
stress on the leaf anatomy of a drought-resistant and a drought-sensitive olive
cultivar. Journal of Horticultural Science & Biotechnology, Vol. 85, No. 4, pp. 289–294,
ISSN 1462-0316
Enstone, D.E.; Peterson, C.A. & Ma, F. (2003). Root endodermis and exodermis : structure,
function, and responses to the environment. Journal of Plant Growth Regulation, Vol.
21, No. 4, pp. 335-351, ISSN 0721-7595
Evans, J.R.; Kaldenhoff, R.; Genty, B. & Terashima, I. (2009). Resistances along the CO2
diffusion pathway inside leaves. Journal of Experimental Botany. Vol. 60, pp. 2235-2248.
Faludi-Daniel, A. (1981). Stomatal behaviour and cuticular properties of maize leaves of
different chilling-resistance during cold treatment. Physiologia Plantarum, Vol.24,
pp. 287-290, ISSN 0031-9317
Fitter, A. & Hay, R. (2002). Environmental Physiology of Plants, Academic Press, ISBN 0-12-
257766-3, San Diego, California
Gong, P.; Zhang, J.; Li, H.; Yang, C.; Zhang, C.; Zhang, X.; Khurram, Z.; Zhang, Y.; Wang, T.;
Fei, Z. & Ye, Z. (2010). Transcriptional profiles of drought-responsive genes in
modulating transcription signal transduction, and biochemical pathways in tomato.
Journal of Experimental Botany, Vol. 61, No. 13, pp. 3563-3575, ISSN 0022-0957
Gottschalk, M.; Dolgener, E.; Xoconostle-Cazares, B.; Lucas, W.J.; Komor, E. & Schobert, C.
(2008). Ricinus communis cyclophilin: functional characterisation of a sieve tube
protein involved in protein folding. Planta, Vol. 228, No. 4, pp. 687-700, ISSN
0032- 0935
Grewal, J.S. & Singh, S.N. (1980). Effect of potassium nutrition on frost damage and yield of
potato plants on alluvial soils of the Punjab (India). Plant Soil, Vol. 57, pp. 105–110
Hare, P.D.; Cress, W.A. & Staden, J.V. (1998). Dissecting the roles of osmolyte accumulation
during stress. Plant, Cell and Environment, Vol. 21, pp. 535–553, ISSN 1365-3040
Hsiao, T.C. & Läuchli, A. (1986). Role of potassium in plant–water relations. In: Advances in
Plant Nutrition, B. Tinker & A. Läuchli (Eds.), Vol. 2, 281-311, Praeger Scientific,
ISBN 0-27592-069-0, New York
IPCC (August 2011). Intergovernmental Panel of Climatic Change WGII, fourth assessment
report, 12/08/2011. Available from http://www.ipcc.ch/publications_and_data/
publications_and_data_reports.htm
Kader, J.C. (1997). Lipid-transfer proteins: a puzzling family of plant proteins. Trends in
Plant Science, Vol. 2, No. 2, pp. 66-70, 1360-1385
Kerstiens, G. (1996). Diffusion of water vapour and gases across cuticles and through
stomatal pores presumed closed. In: Plant cuticles: an integrated functional approach.
G. Kerstiens, (Ed.), 121-134, Bios Scientific Publishers, ISBN 1-85996-130-4,
Lancaster
Kinnersley, A.M. & Turano, F.J. (2000). Gamma aminobutyric acid (GABA) and plant
responses to stress. Critical Reviews in Plant Sciences, Vol. 19, pp. 479–509, ISSN
1549-7836
Kirkham, M.B. (2005). Principles of soil and plant water relations. Elsevier Academic Press,
ISBN 0-12-409751-0, San Diego, California.
Kramer, P.J. & Boyer, J.S. (1995). Water relations of plants and soils. Elsevier Academic Press,
ISBN 0-12-425060-2, San Diego, California
Kutlu N, Terzi R, Tekeli C, Senel G, Battal P & Kadioglu A (2009). Changes in anatomical
structure and levels of endogenous phytohormones during leaf rolling in
Ctenanthe setosa. Turkish Journal of Biology, Vol. 33, pp. 115-122, ISSN 1300-0152
Lafitte, H. R.; Courtois, B. (2002) Interpreting cultivar x environment interactions for yield in
upland rice: assigning value to drought-adaptive traits. Crop Science, Vol. 42, No.
1, pp. 1409-1420, ISSN 0011-183X
Larcher W (1995). Physiological Plant Ecology. Springer-Verlag, ISBN 978-3540435167, Berlin.
Läuchli, A. (1984). Mechanism of nutrient fluxes at membranes of the root surface and their
regulation in the whole plant. In: Roots, Nutrient and Water Influx, and Plant
Growth, S.A. Barber; D.R. Bouldin (Eds.), 1-25, ASA Special Publication, ISBN
0891180826, Madison
Levine, L.H.; Richards, J.T. & Wheeler, R.M. (2009). Super-elevated CO2 interferes with
stomatal response to ABA and night closure in soybean (Glycine max). Journal of
Plant Physiology, Vol.166, No. 9, pp. 903-913, ISSN 0176-1617
Liu X.; Huang B (2000). Heat stress injury in relation to membrane lipid peroxidation in
creeping bentgrass. Crop Science, Vol. 40, Nº1, pp. 503-510.
Maghuly, F.; Borroto-Fernandez, E.G.; Khan, M. A.; Herndl, A.; Marzban, G. & Laimer, M.
(2009). Expression of calmodulin and lipid transfer protein genes in Prunus incisa x
serrula under different stress conditions. Tree Physiology, Vol. 29, No. 3, pp. 437-
444, ISSN 0829-318X
Marschner, H. (1995). Mineral nutrition of higher plants. (2Ed). Academic Press, ISBN 0-12-
4735428, Michigan
matter and root:shoot ratio in cerrado in central Brazil. Brazilian Journal of Ecology, Vol. 2, No.
2, pp. 11-23.
Maurel, C.; Verdoucq, L.; Luu, D. & Santoni, V. (2008). Plant aquaporins: membrane
channels with multiple integrated functions. Annual Review of Plant Biology, Vol. 59,
pp. 595-624, ISSN 0066-4294
Mendgen, K. (1996). Fungal attachment and penetration. In: Plant cuticles: an integrated
functional approach. G. Kerstiens, (Ed.), 175-188, Bios Scientific Publishers, ISBN 1-
85996-130-4, Lancaster
Miglietta, F. & Raschi, A. (1993). Studying the effect to elevated CO2 in the opening a
naturally enriched environment in central Italy. Vegetatio, Vol. 104, No. 105, pp.
391–400, ISSN 0042-3106
Molina-Montenegro, M.A.; Ávila, P.; Hurtado; Valdivia, A.I. & Gianoli, E. (2006). Leaf
trichome density may explain herbivory patterns of Actinote sp. (Lepidoptera:
Archaeidae) on Liabum mandonii (Asteraceae) in a montane humid forest (Nor
Yungas, Bolivia). Acta Oecologica, Vol. 30, pp. 147–150, ISSN 1146-609X
Molinari, H.B.C.,; Marur, C.J.; Bespalhok Filho, J.C.; Kobayashi, A.K.; Pileggi, M.; Leite
Júnior, R.P.; Pereira, L.F.P. & Vieira, L.G.E. (2004). Osmoti adjustment in transgenic
citrus rootstock Carrizo citrange (Citrus sinensis Osb x Poncirus trifoliate L. Raf)
overproducing proline. Plant Science, Vol. 167, pp. 1375–81, ISSN 0168-9452
Montalvo-Hernández, L.; Piedra-Ibarra, E.E.; Gómez-Silva, L.; Lira-Carmona, R.; Acosta-
Gallegos, J.A.; Vazquez-Medrano, J.; Xoconostle-Cázares, B. & Ruíz-Medrano, R.
(2008). Differential accumulation of mRNAs in drought-tolerant and susceptible
common bean cultivars in response to water deficit. The New Phytologist, Vol. 177,
No. 1, pp. 102-113, ISSN 0028-646X
Mullins, G.M.; Bouquet, A.; Williams, L.E. (1992). Biology of the grapevines. Cambridge
University Press, New York. 239 p.
Nobel, P.S. (2009). Physicochemical and environmental plant physiology. (4th Ed). Elsevier, ISBN
978-0-12-374143-1, London
Ogbonnaya, C.I.; Nwalozie, M.C.; Roy-Macauley, H. & Annerose, D.J.M. (1998). Growth and
water relations of Kenaf (Hibiscus cannabinus L.) under water deficit on a sandy
soil. Industrial Crops and Products, Vol. 8, pp. 65–76, ISSN 0926-6690
Poffenroth, M.; Green, D.B. & Tallman, G. (1992). Sugar concentrations in guard cells of Vicia
faba illuminated with red or blue light. Plant Physiology, Vol. 98, pp. 1460–1471,
ISSN 0032-08894
Richter, H. (2001). The cohesion theory debate continues: the pitfalls of cryobiology. Trends
in Plant Science, Vol. 6, No. 10, pp. 456–457, ISSN 1360-1385
Sairam, R.K. & Tyagi, A. (2004). Physiology and molecular biology of salinity stress
tolerance in plants. Current Science, Vol. 86, pp. 407–421, ISSN 0011-3891
Sauter, A.; Dietz, K.J. & Hartung, W. (2002). A possible tress physiological role of abscisic
acid conjugates in root-to-shoot signalling. Plant Cell and Environment, Vol. 25, No.
2, pp. 223-228, ISSN 1365-3040
Scherbatskoy, T. & Tyree, M.T. (1990). Kinetics of exchange of ions between artificial
precipitation and maple leaf surfaces. New Phytologist, Vol. 114, pp. 703-712, ISSN
0028-646X
Scholander, P.F.; Hammel, H.T.; Hemmingsen, E.A. & Bradstreet, E.D. (1965). Hydrostatic
pressure and osmotic potencials in leaves of mangroves and some other plants.
Proceedings of the National Academy Science, Vol. 51, Nº 1, pp.119-125.
Sekhar, K.; Priyanka, B.; Reddy, V.D. & Rao, K.V. (2010). Isolation and characterization of a
pigeonpea cyclophilin (CcCYP) gene, and its over-expression in Arabidopsis
confers multiple abiotic stress tolerance. Plant Cell and Environment, Vol. 33, No. 8,
pp. 1324-1338, ISSN 1365-3040
Seki, M.; Narusaka, M.; Abe, H.; Kasuga, M.; Yamaguchi-Shinozaki, K.; Carninci, P.;
Hayashizaki, Y. & Shinozaki, K. (2001). Monitoring the Expression Pattern of 1300
Arabidopsis Genes under Drought and Cold Stresses by Using a Full-Length cDNA
Microarray. Plant Cell, Vol. 13, No. 1, pp.61-72, ISSN 1040-4651
Shao, H.B.; Chu, L.Y.; Jaleel, C.A. & Zhao, C.X. (2008). Water-deficit stress-induced
anatomical changes in higher plants. Comptes Rendus Biologies, Vol. 331, No. 3,
pp. 215-25, ISSN 1631-0691.
Sharma, S.S. and& K.J. Dietz, 2006. The significance of amino acids and amino acid-derived
molecules in plant responses and adaptation to heavy metal stress. Journal of
Experimental Botany. Vol. 57, pp. 711-726.
Shepherd, T.; Robertson, G.W.; Griffiths, D.W.; Birch, A.N.E. & Duncan, G. (1995). Effects of
environment on the composition of epicuticular wax from kale and swede.
Phytochemistry, Vol. 40, pp. 407-417, ISSN 0031-9422
Sivakumar, P.; Sharmila, P.; Saradhi, P.P. (2000). Proline alleviates salt-stress induced
enhancement in Rubisco oxygenase activity. Biochem Biophys Res Commun, Vol. 279,
Nº 1, pp. 512–5.
Steudle, E. (2001). The cohesion-tension mechanism and the acquisition of water by plant
roots. Annual Review of Plant Physiology and Plant Molecular Biology, Vol. 52, pp. 847–
875, ISSN 1040-2519
Syvertsen, J.P.; Lloyd, J.; Mcconchie, C.; Kriedemann, P.E. & Farquhar, G.D. (1995). On the
relation between leaf anatomy and CO2 diffusion through the mesophyll of
hypostomatous leaves. Plant Cell and Environment, Vol. 18, pp. 149–157, ISSN
1365-3040
Taiz, L. & Zeiger, E. (2002). Plant physiology. (3rd Ed.). Sinauer Associates, ISBN 0-87893-823-
0, Sunderland.
Treviño, M.B. & O'Connell, M.A. (1998). Three drought-responsive members of the
nonspecific lipid-transfer protein gene family in Lycopersicon pennellii show
different developmental patterns of expression. Plant Physiology, Vol. 116, No. 4,
pp. 1461-1468, ISSN 0032-0889
Tyree, M.T. & Sperry, J.S. (1989). Vulnerability of xylem to cavitation and embolism.
Annual Review of Plant Physiology and Plant Molecular Biology, Vol. 40, pp. 19–38,
ISSN 1040-2519
Tyree, M.T. & Zimmermann, M.H. (2002). Xylem Structure and the Ascent of Sap. (2Ed).
Springer series in wood science, Springer-Verlag, ISBN 3-540-43354-6, Berlin
Tyree, M.T. (2003). Plant hydraulics: the ascent of water. Nature, Vol. 423, No. 6943, pp. 923-
923, ISSN 0028-0836
Vanrensburg, L.; Kruger, G.H.J.; Kruger, R.H. (1993). Proline accumulation the drought
tolerance selection: its relantionship to membrane integrity and chloroplast ultra
structure in Nicotiana tabacum L. Journal of Plant Physiology, Vol. 141, Nº1, pp.188–94.
Vigh, L.; Horváth, I.; Farkas, T.; Mustardy, L.A. & Vogelmann, T.C. (1993). Plant-tissue
optics. Annual Review of Plant Physiology and Plant Molecular Biology, Vol. 44, pp.
231–251, ISSN 1040-2519
Vogelmann, T.C. (1993). Plant tissue optics. Annual Review Plant Physiological and Plant
Molecular biology. Vol. 44, pp. 231-251.
Wang, W.X.; Vinocur, B.; Shoseyov, O. & Altman, A. (2004). Role of plant heat-shock
proteins and molecular chaperones in the abiotic stress response. Trends in Plant
Science, Vol. 9, No. 5, pp. 244-252, ISSN 1360-1385
Xie, X.; Wang, Y.; Williamson, L.; Holroyd, G.H.; Tagliavia, C.; Murchie, E.; Teobald, J.;
Knight, M.R.; Davies, W.J.; Leyser, H.M. & Hetherington, A.M. (2006). The
identification of genes involved in the stomatal response to reduced atmospheric
relative humidity. Current Biology, Vol. 16, No. 9, pp. 882-887, ISSN 0960-9822
Xoconostle-Cázares, B.; Ramírez-Ortega, F.A.; Flores-Elenes, L. & Ruiz-Medrano, R. (2010).
Drought tolerance in crop plants. American Journal of Plant Physiology, Vol. 5, No. 5,
pp. 241-256, ISSN 0176-1617
Yokota, A.; Takahara, K & Akashi, K. (2006). Water Stress. In: Physiology and Molecular
Biology of Stress Tolerance in Plants, K.V. M. Rao; A.S. Raghavendra & K.J. Reddy
(Eds.), 15-39, Springer, ISBN-10 1-4020-4224-8, Netherlands
Zhu, X.; Gong, H.; Chen, G.; Wang, S. & Zhang, C. (2005). Different solute levels in two spring
wheat cultivars induced by progressive field water stress at different developmental
stages. Journal of Arid Environments, Vol. 62, pp. 1–14, ISSN 0140-1963
Zimmermann, U.; Schneider, H; Wegner, L.H. & Haase, A. (2004). Water ascent in tall trees:
does evolution of land plants rely on a highly metastable state? New Phytologist,
Vol. 162, No. 3, pp. 575–615, ISSN 1469-8137
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Plant Water Relations: Absorption, Transport and Control

Chapter · April 2012

Geraldo Chavarria Henrique Pessoa dos Santos

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Plant Water Relations: Absorption,

2. Absorption and water flow through plants

2.1 Water potential

2.2 Water dynamics in soil-plant-atmosphere system

2.2.1 Soil water

2.2.2 Water absorption by the roots

demands of photo-transpiration. The aquaporins are controlled by many endogenous and

2.2.3 Ascension of water through the plant: Vascular system

Jv = (πR4∆Ψ) / 8ηL (1)

2.2.4 Leaf water and transpiration

Water in cell wall

air layer limitrophe Water

3. Physiological and biochemical aspects of water in plants

3.1 Water deficit and its effects on plant growth

3.2 Elements that define water demand

3.3 Mechanisms of water status regulation

3.3.2 Stomata metabolism

Guard cell turgid

Guard cell flacid

3.3 Hormonal and molecular responses in different water conditions

3.4 Osmotic regulation

4. Practical aspects of measuring plant water status

Ainsworth, E. A. & Rogers, A. (2007). The response of photosynthesis and stomatal

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