Set 1--Answers to selected problems

Water potential

3. A cell with a pressure potential of 0.8 MPa and an osmotic

potential of -1.6 MPa is placed in a beaker of pure water. What is the
water potential in the beaker and in the cell initially? In which
direction will water flow? After equilibrium, what will be the
component potentials and water potentials in the beaker and cell?
What assumptions have you made in your calculations?

Initially, Ψw of the cell is -0.8 MPa, and Ψw of the water is 0. Water

flows from outside to inside the cell. At equilibrium, Ψw of the cell is
0, Ψs = -1.6, Ψp = 1.6 MPa, assuming that the increase in cell volume
did not significantly dilute the cell contents. If you wish, you can
assume that the cell volume did increase, say 10%, and the content
decreased to 90% of the initial value. In that case,

Ψs = -1.44 (from -1.6 x 90%) and Ψp = 1.44 MPa.

4. If the cell in the previous questions is placed in a beaker that

contains a solution with an osmotic potential of -0.7 MPa , which
direction will water flow? Compute your answer both assuming the
cell is used before and after equilibrium in water. What will be the
values of the water potential , pressure potential, and osmotic
potential in the cell at equilibrium?

If the cell is placed in the beaker before being in water, water flows
into the cell; if after being in water, water flows out. At equilibrium,
Ψw of the cell is -0.7, Ψp = 0.9, Ψs = -1.6 MPa (again, assuming no
significant increase in volume).

7. The primary walls of leaf mesophyl cells in some plants are much
more elastic (that is, stretchy) than in other plants. For example, a
willow, Salix lasiandra, which grows near streams, has more elastic
cell walls than Ginkgo biloba trees, which grow in drier areas. (a) If I
gave you a corn leaf and a sunflower leaf, how would you tell how
elastic the cells of each were? (b) What adaptive advantage could
greater elasticity (stretchiness) provide? Hint: consider that growth
rate is related to Ψp. (c) What disadvantage might be associated with
greater elasticity?

(a) Apply different forces and measure extension; plot force vs

extension (relative extension, Δl/l); lower slope, that is, less force per
unit extension, is more stretchy. (b) If growth rate really is related to
extension, then greater stretchiness would allow greater growth at a
given force. (c) Greater stretchiness might disrupt more connections
between cells, if adjacent cells stretched different amounts.

8. The eggs of the marine alga Fucus are shed directly into sea water
without any cell walls. This makes it easier for the antherozoids
(sperm) to fertilize them. A cell wall forms 4 to 8 hours after
fertilization. About 15-25 hours after fertilization, the cell starts to
enlarge, forming a long, thin rhizoid. Discuss the components of
water potential of the Fucus egg before fertilization, immediately
after wall formation, and during the formation of the rhizoid. Assume
the seawater has a Ψs of -0.7 MPa.

Before fertilization, Ψw = -0.7 MPa, Ψp = 0, Ψs = -0.7 MPa, and this

is probably true immediately after wall formation, also. During
formation of the rhizoid, Ψp > 0 (pressure is needed to expand the
wall), but still, Ψw = -0.7 MPa, so Ψs < -0.7 MPa.

10. A fresh lettuce leaf with a 90% relative water content is placed in
a box at 20oC that contains air of 90% relative humidity. Will the leaf
lose water, absorb water, or stay the same? If the box is put in a
refrigerator at 4oC, will the leaf lose water, absorb water, or stay the
same?

A "fresh" lettuce leaf can be assumed to be turgid and thus have Ψp>
0 and Ψw > -0.7 MPa. Air at 90% RH has Ψ = -14.3 MPa. The leaf
will lose water to air. Placed into a refrigerator, the air in the box will
become saturated (100% RH), Ψ = 0 MPa, and the leaf will gain
water.

11. Calculate the original Ψcell, Ψs, and Ψp for the cells in a thin slice
of plant tissue, using data that show the gain and loss of weight, and
the % plasmolysis, when the tissue is placed in different
concentrations of sucrose. What errors are inherent in this method?

Sucrose concentration Weight %

(Molal) change (g) plasmolysis

0 + 0.53 0

0.05 +0.50 0

0.1 +0.40 0

0.15 +0.15 0

0.2 -0.05 0

0.25 -0.20 10

0.30 -0.40 90

Start by plotting the weight change and fraction of plasmolyzed cells

by the water potential [-2.44 x sucrose concentration (MPa)]. The
abscissa point where weight change is 0 is the original Ψcell of the
tissue. The point of incipient plasmolysis (estimated point at which
no cells are plasmolyzed) is where Ψp = 0 and the abscissa indicates
Ψs. Assuming that Ψs is about the same under the original
conditions, you can calculate the original Ψp as Ψcell - Ψs. Ψcell = -0.48
MPa; Ψs = -0.58 MPa, Ψs = 0.1 MPa. The calculations assume that
the tissue is homogeneous, that the tissue is at equilibrium when the
observations are made, and that the tissue has not changed during the
assay.

Transpiration

13. A small branch is cut from a cottonwood tree on an early spring

morning, when the temperature is 10oC and there is a light fog. The
branch is placed in a Scholander pressure bomb. It takes 1.0 MPa of
pressure to push the xylem sap to the end of the branch so that it is
just visible. (a) Is the system in the bomb at equilibrium with regard
to water potential? (b) Assuming that Ψs(xylem) = 0, calculate
Ψ(xylem) and Ψ(leaf cell) when the branch was on the tree and
when the branch is in the bomb. (c) What differences would you
expect if the experiment were repeated later in the day, when the
temperature was 30oC, the fog had disappeared, and there was a brisk
wind?

(a) The system inside the bomb is at equilibrium. (b) In the bomb,
Ψw(xylem) = 0 (it is at equilibrium with air outside the bomb), and
Ψw(leaf cell) = 0; on the tree, Ψw(xylem) = -1 MPa, and Ψw(leaf cell)
= -1 MPa. (c) The plant will be transpiring, and both Ψw(xylem) and
Ψw(leaf cell) would be < -1 MPa. On the plant (not in the bomb),
Ψw(leaf cell) < Ψw(xylem), the amount of difference depending on
the transpiration rate. On the plant, the system is not at equilibrium.

15. Water has certain physiochemical properties; for example, strong

cohesive forces exist between individual water molecules, and one
result of these forces is that water has a high surface tension. Explain
why these two properties are so important in terms of water
movement in the plant.

Cohesion prevents water columns in xylem from breaking. Surface

tension prevents air from entering xylem through pits, keeps cells
walls wet, and increases the water capacity of fine-grained soils.

19. Would a tub of water maintained at 30oC lose water by

evaporation more rapidly on a cold, clear winter day (temperature
0oC) or on a warm summer day (temperature 35oC)? Assume 40%
(0.4) relative humidity at both temperatures.

The rate of water loss will depend directly on dc/dx (Δc/Δx,

considering a linear gradient across an unstirred boundary layer).
Assume Δx is the same for both tubs. Δc will be the difference
between c just above the tub (saturated at 30oC, 1.687 mol/m3) and c
in bulk air, 0.4 x 0.269 = 0.108 at 0oC and 0.4 x 2.201 = 0.880 at
35oC. The difference is greater at 0oC.

22. Provide a detailed description of why plants can use the energy
from the sun to drive the flow of water through the plant.

Under most circumstances, water is pulled through the plant by the

"tug", exerted when water evaporates due to negative water potential
of the air. The energy of the sun warms the leaf and the air in the leaf,
which increases its saturation vapor concentration, which in turn
increases the gradient between the vapor concentration in leaf and
the outside air. More indirectly, water can be pushed into the xylem
by "root pressure," which comes from accumulation of solutes in the
xylem fluid. The energy of the sun is involved through
photosynthesis, phloem transport, and respiration to power the
accumulation.

Energy balance and water stress

27. An article by Boyer (Science 218:773, 1982) demonstrated how

plant breeding could increase crop yield. His point was that new
varieties were better at extracting water from the soil and transferring
it to the leaves. Discuss the genetic traits that might have led to
increased water uptake ability. What other traits (besides those
leading to increased water uptake ability) might affect average
afternoon water potential of the leaves?

Increased water uptake could be due to: more root branching, more
root per shoot, more aquaporins in root cell membranes (reducing
resistance), larger tracheid and vessel diameters (reducing resistance),
more vessels compared to tracheids, more xylem in stems, petioles,
and leaves. Other possible traits include: control of stomatal aperture
in response to water stress or time-of-day.

29. Water in the plant can be described as a system that is constantly

experiencing a "tug-of-war." Using the special properties of water
and the anatomical features of the plant, explain this "tug-of-war"
phenomenon.

"Tug" at the top (water-air interface in the leaf) is due to the force
exerted by surface tension in small capillaries (at the pits of xylem, in
the cell walls of parenchymal cells). Another source of "tug"--inside
the cell--is provided by the solute effect. Tug at the bottom of the
xylem is provided by gravity, by solute effect of stelar and cortical
cells, and by matric potential of soil (also surface tension). The
balance requires cohesion of water through intermolecular hydrogen
bonding.

34. A small bean plant is growing under a bell jar in a saturated

atmosphere at 25oC. As the sunlight strikes the leaf, its temperature
rises 10oC above the air temperature. A similar bean plant is growing
under a cloth shade nearby. Air temperature is 25oC, relative
humidity is 70%, and leaf temperature is 25oC. Which plant can be
expected to have the more rapid transpiration rate? Compute the
rates, assuming Rs = 100 s m-1 and Ra = 50 s m-1.

The plant in the bell jar transpires more rapidly. Bell jar: Δc = 2.201
- 1.28 = 0.921 mol/m3; rate = 20000*0.921/150 = 122.8 mg/m2-s.
Shade: Δc = 1.28 - 0.7(1.28) = 0.384 mol/m3; rate =
20000*0.384/150 = 51.2 mg/m2-s.

36. Some crop plants tend to grow mostly at night, while others grow
more or less uniformly day and night. Discuss differences in the
plants that could account for the differences in growth patterns,
considering especially net water potential and stomatal control.

Growth depends on Ψp and wall structure (yield threshold and

response to pressure above the yield point). Plants that grow mostly
at night are responding to higher Ψw and Ψp. Plants that grow
uniformly must (a) be well watered (uniform Ψw), (b) close stomata
to maintain a high Ψw, (c) adjust Ψs to keep Ψp constant, and/or (d)
adjust their wall structure to vary the yield threshold or response.

37. A small herbaceous plant growing on a sparsely covered sand

dune has narrow, leathery leaves, which are green on top but almost
pure white underneath. Closer inspection shows a thick mat of white
hairs on the bottom surface. What effect will these hairs have on the
plant’s mid-day energy balance, if (a) all the stomata are on the top
surface; or if (b) all the stomata are on the bottom surface?

Leaf hairs have two effects: reflecting light/IR radiation and

stabilizing the boundary layer. In (a), they do not affect the boundary
layer and transpiration, so their effect is to reduce +Q, with a
generally cooling effect. In (b), they still reduce +Q, but they also
may raise (make less negative) -V. The net effect depends on the
relative values of +Q (from below) and -V.

41. Oranges developing in orchards in winter are very sensitive to

freezing. The problem can be countered when it occurs on clear,
calm nights. There are several techniques. Smudgepots (burning oil,
which produces thick smoke) were used in the past, but no longer
because of the pollution they cause. Sprinklers are popular, as are
large fans. How does each help the problem?

The problem occurs on clear nights, because there is very little

incoming Q from space (or air). The trees and ground radiate Q
outward, and their temperature, and the temperature of the surface air,
drops below freezing. The cold air, being denser, stays at the surface.
Smudgepots produce smoke, which radiates Q to the ground (and
absorbs and re-irradiates the Q from the trees and ground).
Sprinklers, which increase the amount of water vapor in the air, do
the same thing. In addition, water that freezes releases heat. Large
fans remove the cold layer of air and warm the trees by convection
with the warmer air from above.

42. Tobacco plants, like most plants, open their stomata during the
day and close them at night. Kalenchoe, a so-called "CAM plant,"
opens stomata during the night and closes them during the day.
What differences would you expect between the guard cells of the
two different types of plants?

In CAM plants, Ψp becomes more positive, because Ψs becomes

more negative, at night, rather than in the day. Either the blue-light
photoreceptor or circadian control, or both, at night stimulate the
conversion of starch to malate, which serves as counter-ion for
accumulated K+. The control is, at least in part, manifested in the
phosphorylation and activation of PEP carboxylase. In C3 plants,
this activation occurs in the light. (In CAM plants, malate is also
formed as a CO2 storage form in mesophyll cells.)

45. Explain the following observation: A row of beans and a row of

pumpkin plants are adjacent in a Davis garden. During the course of
each day, the pumpkin leaves wilt, but the beans leaves do not.

We can assume that this is temporary wilt and thus that Ψw of soil is
less negative than Ψs of leaf cells (Ψp of leaf cells is positive at night).
The wilt indicates that the Ψw of the leaf apoplast falls below Ψs of
the leaf cells during the day. This must occur in pumpkin but not
bean, either because Ψs of pumpkin leaf cells is less negative than
that of bean, or because of resistance to water flow up the pumpkin
xylem or into the root stele is greater than in bean. If the Ψs of the
bean cells is more negative than that of pumpkin cells, it may be so
all the time, or it may adjust daily as Ψw of the soil falls.

46. List several ways in which water deficit can affect shoot growth.
Which is permanent in effect even when sufficient water
subsequently becomes available?

Reduce Ψp of cells below yield threshold; change wall synthesis

(increasing yield threshold--permanent); inhibit metabolism (protein
synthesis, cell division--permanent); induce ABA accumulation;
close stomata (inhibiting photosynthesis).

52. Using the Zea mays stomatal guard cell system as your model,
explain the events that may give rise to stomatal closure during a
period of water stress. Start with “Ψw of soil drops.”

Ψw of soil drops -> Ψw and Ψp of root cells drop -> ABA synthesis -
> ABA exported to xylem sap and moves to leaf in transpiration
stream [and] xylem sap pH rises from 6.3 to 7.2 (why? how?) ->
ABA moves to guard cells (receptors in plasma membrane or in
cytoplasm--see p. 549 ff) -> increased cytoplasmic Ca2+, inhibition of
H + efflux, opening of anion channels -> efflux of Cl- and malate- ->
depolarization of membrane potential -> K+ efflux -> less negative Ψs
-> less positive Ψp -> elastic contraction of cell walls -> stoma closes

Mineral Nutrition

54. Use the molecular definition of an essential element to discuss

the roles played by Mg2+ in the physiological process of a generic
plant. Consider how the plant might respond to changes in Mg2+
levels in the soil in terms of supplying Mg2+ to the body of the plant.

Molecular definition: the element is required for the structure or

biochemical activity of an component that is essential for life. Mg2+ is
essential for many enzymes involving DNA or RNA synthesis, ATP
use, phosphate transfer; constituent of chlorophyll. (In contrast, the
physiological definition is an element required for optimum growth,
development, and reproduction.) In a situation where Mg2+
concentration is low, a plant might (a) induce carriers with greater
affinity (if genes for such carriers are available); (b) mobilize Mg2+
from the lower, mature leaves to support further growth of roots and
flowers.

55. Describe and explain the differences between the nutritional

requirements of a plant and an animal.

Plant requirements are all inorganic, including some oxidized ions

that animals cannot reduce, like NO3- and Fe3+. Energy is obtained
from light (although in culture, plants can obtain energy by respiring
supplied carbohydrates, e.g. sucrose). Animals have some essential
organic nutrients (humans: amino acids, some fatty acids, vitamins).

58. Discuss the principle of an essential mineral nutrient with respect

to the cations Ca2+ and Fe3+, and the anion NO3-.

Ca2+ is essential for cell wall and membrane structures and for the
activity of some enzymes, and it serves as a signal compound; a lack
results in necrosis of growing regions. Fe3+ must be reduced to Fe2+,
which is essential for chlorophyll synthesis and for some cofactors
(cytochromes) in mitochondrial respiratory metabolism; a lack results
in chlorotic young leaves. NO3- is a source of N, essential for the
synthesis of any amino acid and nucleotide, but it must be reduced to
NH4+ before it can be incorporated into glutamine and from there to
other compounds; a lack results in chlorosis of older leaves, stunting,
and brittleness from excess carbohydrates.

59. Boron has been recognized as an essential element only recently.

Explain why it was so difficult to make this determination.
The concentration that is toxic is only 2-3-fold higher than the
concentration below which one gets deficiency symptoms. This
means that it is difficult to find an optimum level. That observation,
coupled with the fact that B is a contaminant in some chemicals that
are needed to make a nutrient solution for hydroponic plant growth,
made it difficult to show that adding B had a promotive effect.