Lecture 12

Resource acquisition and water transport

I. Introduction
A. Plants have various ways by which they can maximize their photosynthetic efficiency.
1. Stem provides conduit by which water and minerals, and food transport may
occur.
- Water and minerals are transported from the roots up into the stem and
finally to leaves. Food is transported to anywhere in the plant that requires
it.
2. Phyllotaxy is the arrangement of leaves on stem.
- One reason for the variations described below is to minimize the shading
of leaves in order to maximize photosynthesis.
a. Alternate or spiral arrangement. Alternate arrangement has leaves
arranged on alternate sides of the stem, usually spiraling around stem. In
either case, there is one leaf per node.
b. Opposite leaf arrangement has two leaves on opposite side of the same
nodes, with succeeding pairs of leaves being at right angle to the
previous pair of leaves.
c. Whorled leaf arrangement has three or more leaves on the same node.
d. Regardless of how leaves are arranged, there is an optimal number of
leaves that is not exceeded to minimize shading.
3. Leaf surface exposed to sunlight.
a. Under low-light conditions, leaves are borne horizontally to maximize
photosynthesis.
i. In dark understory, as found in tropical rainforest, leaves with large
surface areas evolved to better absorb what little light is available.
b. Under high light conditions, vertical leaves will protect lower leaves
against high intensity sunlight and expose less surface areas of their
leaves.
4. Height of tree and branching patterns also important.
c. Many palms are tall and unbranched, with leaves at the very top to
maximize photosynthesis.
d. Many eudicots are shorter, but with lots of branching, with smaller
leaves that do not overlap each other.
B. Root architecture and acquisition of water and minerals.
1. Woody eudicots and gymnosperms with long taproots and lateral roots more
 readily access water and minerals and for this reason are bigger trees than
monocots.
2. Monocots have fibrous roots that are shallow and access to water and mineral
not as great as in above. Reason why their trees are usually shorter.
3. Short distant transport of water and minerals in roots to endodermis.
a. Movement through two major pathways:
i. Apoplast: Non-living components of cell, i.e., cell wall and
intercellular space.
ii. Symplast: Living components of cell, i.e., protoplast and
plasmodesmata.
iii. Transmembrane: Both apoplast and symplast occurs.
b. Can only be symplast once endodermis is reached!
i. Endodermis has casparian strip on radial wall that is impermeable to
water. This allows selection as to what ions are allowed to go
through.
ii. Once water and minerals are beyond endodermis, they cannot “leak”
back out and must enter vascular tissue.
5. Once in vascular tissue, another mechanism takes over for water and mineral
transport that is a long distant transport mechanism.
c. Root pressure is a “pushing” mechanism. Although discussed in your
text, it is not a very credible mechanism for long distant transport of
xylem sap for reason discussed in lecture. Does not occur in all plants
and not in gymnosperms at all.
d. Cohesion-Tension Hypothesis is a “pulling” mechanism and is believed
to be long distant transport of water.
6. Mechanism for cohesion-tension hypothesis.
a. Transpiration is driving force. Evaporation of water through stomata.
i. Transpiration creates negative pressure that pulls xylem sap up xylem.
This long distant transport is called bulk flow and is
independent of osmosis and active transport.
b. Transpiration pull is important, but property of water is also important in
order for mechanism to work.
i. Water molecule is polar and is held together by hydrogen bonding,
i.e., cohesion of water molecules. If hydrogen bond breaks,
cavitation occurs and bubble forms in vessel element or tracheid.
Water above bubble still can be transported to leaves, but water
below bubble, can no longer go up this particular vessel elements.
However, lateral transport through pits, and then up adjacent cells
 can occur.
ii. Adhesion of water molecules to cell wall of vessel element also
important in support of water molecules in xylem.
iii. Surface tension of water molecule at interface of mesophyll cells
where water is constantly being evaporated. As surface tension
increases, negative pressure in pulling evaporating water out of leaf,
through stomata.
iv. Most water is lost through stomata, but enough is utilized in
photosynthesis to make food for plant. However, a balance of water
loss and photosynthesis must occur.
7. Stomata regulate transpiration and maximize photosynthesis. b. Stomata
formed by pair of guard cells that become turgid, when open and flaccid,
when closed.
i. Radial microfibrils on guard cell and increase in turgor causes
“bending” of the guard cells, forming the opening of stoma.
ii. Increase in turgor comes about through K+ transport in and out of
guard cells.
iii. Important that not too much water lost through evaporation so that
plant remains turgid, but this is at a cost to photosynthesis, when
transpiration must be slowed in order not to lose too much water.
8. Change in environment effects rate of transpiration:
b. Humidity, when high outside of plant slows transpiration and when low
increases transpiration.
c. High air temperature increases transpiration. Low temperature decreases
it.
d. Insufficient water in soil, closes stomata and decreases transpiration rate.
9. Plant adaptations and transpiration.
b. Plants evolved in very dry environments have evolved various
mechanisms to decrease transpiration.
i. Crassulacean acid metabolism (CAM). Stomatal gas exchange
occurs at night to prevent water loss.
ii. Cactus plants have modified leaves that have evolved into spines that
reduce transpiration.
iii. Plants with several epidermal layers on upper surface of leaves that
do not have stomata. Stomata only on lower epidermis.
iv. Stomata that form inside the “crypts” of leaves that have trichomes
(=leaf hairs) that will hold free water. High humidity in crypts
reduce transpiration.
 v. Plants that dormant and are stimulated to grow leaves, flower and set
seed during a short rainy season.
c. Plants evolved in very wet and hot environments, e.g., tropical rain
forest.
i. Crowded conditions favor tall trees with leaves higher up in order to
not get shaded by adjacent trees. Resulting in dark canopy below.
ii. Plants on forest floors have large leaves in order to capture more
light, in the dimly lit under story. Loss of water not a problem with
increased transpiration since sufficient water not a problem. Very
high rate of respiration.

Terms to know:

Phyllotaxy- the arrangement of leaves on stem

Alternate or spiral arrangement of leaves - leaves arranged on alternate sides of the stem, usually
spiraling around stem, one leaf
Opposite arrangement of leaves
Whorled arrangement of leaves
Apoplast
Symplast
Transmembrane
Casparian strip
Endodermis
Root pressure
Cohesion-tension hypothesis
Polar molecule
Hydrogen bonding
Cohesion (of water molecules)
Adhesion (of water molecules to cell wall.)
Transpiration
Negative pressure
Bulk flow
Surface tension
Cavitation

Questions

1. What is phyllotaxy?
The arrangement of leaves on stem
2. What are different ways in which leaves may be organized on plants? What is one
possible reason for the variations in which leaves are arranged on plants.

3. What are the advantages of tall trees with little or not branching and short trees with lots
of branches? Each have their advantages in their proper environments. What is the
advantage of each?

4. What is the significance (function) of the endodermis in short distant transport of water
and minerals?

5. How does root pressure transport water in plants?

6. What are the key concepts in understanding long distant transport of xylem sap, i.e.,
transpiration, bulk flow, transpiration, cohesion and adhesion of water molecules,
tension and cavitation?

7. What happens to the water in vessel elements and tracheids in which cavitation has
occurred?

8. The opening and closing of stomata on the leaves impact the rate at which transpiration
occurs. Explain how the guard cells open and close stomata and how this impacts the rate
of transpiration.

9. What are some key adaptations that desert plants have evolved in order to reduce their
rates of transpiration, in order to reduce water loss while still maximizing
photosynthesis?

10. What are some key adaptations that tropical plants have evolved that enable them to
maximize photosynthesis? Loss of water in transpiration is not that much of an issue in
 this environment, but transpiration can often be decreased. Why?

Hydrenchyma cells function in storage of excess water.