Absorption of Water
Terrestrial
plants have to absorb water from the soil to maintain turgidity, metabolic
activities and growth. Absorption of water from soil takes place in two steps:
1.
From soil to root hairs – either actively or
passively.
2.
From root hairs further transport in the lateral
direction to reach xylem, the superhighway of water transport.
Usually,
absorption of water occurs in plants through young roots. The zone of rapid
water absorption is root hairs. They
are delicate structures which get continuously replaced by new ones. Root hairs
are unicellular extensions of epidermal cells without cuticle. Root hairs are
extremely thin and numerous and they provide a large surface area for
absorption (Figure 11.10).
Water is
first absorbed by root hair and other epidermal cells through imbibition from soil
and moves radially and centripetally across the cortex, endodermis, pericycle
and finally reaches xylem elements osmotically.
There are
three possible routes of water (Figure 11.11). They are i) Apoplast ii) Symplast iii)
Transmembrane route.
The apoplast (Greek: apo = away; plast
= cell) consists of everything external
The apoplast includes cell walls, extra
cellular spaces and the interior of dead cells such as vessel elements and
tracheids. In the apoplast pathway, water moves exclusively through the cell
wall or the non-living part of the plant without crossing any membrane. The
apoplast is a continuous system.
The symplast (Greek: sym = within; plast
= cell) consists of the entire mass of cytosol of all the living cells in a
plant, as well as the plasmodesmata,
the cytoplasmic channel that
interconnects them.
In the
symplastic route, water has to cross plasma membrane to enter the cytoplasm of
outer root cell; then it will move within adjoining cytoplasm through
plasmodesmata around the vacuoles without the necessity to cross more membrane,
till it reaches xylem.
In
transmembrane pathway water sequentially enters a cell on one side and exits
from the cell on the other side. In this pathway, water crosses at least two
membranes for each cell. Transport across the tonoplast is also involved.
Kramer (1949) recognized two distinct mechanisms which independently operate
in the absorption of water in plants. They are, i) active absorption ii)
passive absorption.
The
mechanism of water absorption due to forces generated in the root itself is
called active absorption. Active
absorption may be osmotic or
non-osmotic.
The
theory of osmotic active absorption was postulated by Atkins (1916) and Preistley
(1923). According to this theory, the first step in the absorption is soil
water imbibed by cell wall of the root hair followed by osmosis. The soil water
is hypotonic and cell sap is hypertonic. Therefore, soil water diffuses into
root hair along the concentration gradient (endosmosis). When the root hair
becomes fully turgid, it becomes hypotonic and water moves osmotically to the
outer most cortical cell. In the same way, water enters into inner cortex,
endodermis, pericycle and finally reaches protoxylem. As the
sap
reaches the protoxylem a pressure is developed known as root pressure. This theory involves the symplastic movement of
water.
Objections to osmotic theory:
1.The cell sap concentration in xylem is not always high.
2. Root pressure is not universal in all plants especially in
trees.
Bennet-Clark (1936), Thimann (1951) and Kramer (1959) observed absorption of
water even if the concentration of cell sap in the root hair is lower than that
of the soil water. Such a movement requires an expenditure of energy released
by respiration (ATP). Thus, there is a link between water absorption and respiration.
It is evident from the fact that when respiratory inhibitors like KCN,
Chloroform are applied there is a decrease in the rate of respiration and also
the rate of absorption of water.
In
passive absorption, roots do not play any role in the absorption of water and
is regulated by transpiration only. Due to transpiration, water is lost from
leaf cells along with a drop in turgor pressure. It increases DPD in leaf cells
and leads to withdrawal of water from adjacent xylem cells. In xylem, a tension
is developed and is transmitted downward up to root resulting in the absorption
of water from the soil.
In
passive absorption (Table 11.3), the path of water may be symplastic or
apoplastic. It accounts for about 98% of the total water uptake by plants.
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