Homeostasis and Adaptation

36

Water Balance in Plants

Without sufficient water plant cells will lose turgor and the plant
tissue will wilt. If the plant passes its permanent wilting point the
plant will die. Water is lost from the plant by transpiration: the loss
of water vapour, primarily through the stomata. Water balance IS not
a problem for aquatic plants. They simply allo.w water to flow 1n by
osmosis until the cell wall stops further expans1on. Plants adapted to

Tropical Forest Plant

leaves modified into

spines or hairs to
reduce water loss

Ocean Margin Plant

Surface area reduced

by producing a squat,
rounded plant shape

Mangrove trees take in brackish

water, excreting the salt through
glands in the leaves

Desert plants e.g. cacti, cope with low rainfall

and high transpiration rates. Plants develop
strategies to reduce water loss, store water,
and access available water supplies.

TS of Marram grass leaf

Ice plant (Carpobrotus): The leaves of many desert and beach dwelling
plants are fleshy or succulent. The leaves are triangular in cross section
and crammed with water storage cells. The water is stored after rain for
use in dry periods. The shallow root system is able to take up water from
the soil surface, taking advantage of any overnight condensation.

Stem becomes the

major photosynthetic
organ, plus a reservoir
for water storage

Marram grass (Ammophi/a): The long, wiry leaf blades of

this beach grass are curled downwards with the stomata on
the inside. This protects them against d rying out by providing
a moist microclimate around the stomata. Plants adapted to
high altitude often have similar adaptations.

land plants that colonise the shoreline (e.g.

mangroves) must cope with high salt
content in the water. Seaweeds below low
tide do not have a water balance problem.

Grasses living in dry areas curl their Mosses are poor at obtaining and Hairs on leaves trap air close to the Excess water is forced from leaves
leaves and have sunken stomata.
storing water, restricting distribution. surface, reducing transpiration rate. (guttation) during high humidity.

Effect of Adaptation

Ball cactus (De/osperma saturatum): In cacti, the leaves are modified into long, thin spines which project outward from the thick fleshy stem
(see close-up above right). This reduces the surface area over which water loss can occur. The stem takes over the role of producing the
food for the plant and also stores water during rainy periods for use during drought. As in succulents like ice plant, the root system in cacti
is shallow to take advantage of surface water appearing as a result of overnight condensation.

1. Define the term xeromorphic: - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

Methods of ~afer Conservation in Various Plant 'Species

Adaptation for Water Conservation

l eaf hairs

Seaweeds growing in the

intertidal zone tolerate exposure
to the drying air every 12 hrs

Shallow, but
extensive
fibrous root
system

Tropical plants live in areas of often high

rainfall. There is also a corresponding high
transpiration rate. Water availability is not a
problem in this environment.

Adaptations in Halophytes and Drought Tolerant Plants

low water conditions are called xerophytes and they exhibit

structural (xeromorphic) and physiological adaptations for water
conservation. Some of these are outlined below. Halophytes (salt
tolerant plants) and alpine species may also show xeromorphic
features: an adaptation to the scarcity of physiologically available
water and high transpirational losses in these environments.

Dry Desert Plant

Rain is channelled by
funnel shaped leaves

37

Homeostasis and Adaptation

Example

2. Describe three xeromorphic adaptations of plants:

Thick, waxy cuticle to stems and leaves

Reduces water loss through the cuticle.

Pinus sp. ivy (Hedera), sea holly

(Eryngium), prickly pear (Opuntia)

Reduced number of stomata

Reduces the number of pores through

which water loss can occur.

Prickly pear (Opuntia), Nerium sp.

Stomata sunken in pits, grooves, or depressions

l eaf surface covered with fine hairs
Massing of leaves into a rosette at ground level

Moist air is trapped close to the area of

water loss, reducing the diffusion gradient
and therefore the rate of water loss.

Sunken stomata: Pinus sp., Hakea sp.;

Hairy leaves: l amb's ear; l eaf rosettes:
Dandelion (Taraxacum), daisy

Stomata closed during the light, open at night

Carbon dioxide is fixed during the night,

water loss during the day is minimised.

CAM plants e.g. American aloe,

pineapple, Kalanchoe, Yucca

leaves reduced to scales, stem photosynthetic

leaves curled, rolled, or folded when flaccid

Reduction in surface area from which

transpiration can occur.

leaf scales: Broom (Cytisus); Rolled leaf:

Marram grass (Ammophila), Erica sp.

Fleshy or succulent stems

Fleshy or succulent leaves

When readily available, water is stored in

the tissues for times of low availability.

Fleshy stems: Opuntia, Candle plant

(Kieinia); Fleshy leaves: Bryophyllum

Deep root system below the water table

Roots tap into the lower water table.

Acacias, oleander

Shallow root system absorbing surface moisture

Roots absorb overnight condensation.

Most cacti

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Biozone International 2000

(a) _________________________________________________________________
(b) _ _ _ __ __ _ _ _ __

_ __ _ _ _ _ _ _ __ _ __ _ _ _ ___

(c) __________________________________________________________________

3. Describe a physiological mechanism by which plants can reduce water loss during the daylight hours:

4. Explain why creating a moist microenvironment around the areas of water loss reduces transpiration rate:

5. Explain why seashore plants (halophytes) exhibit many desert-dwelling adaptations: _ _ _ _ _ _ _ __ _ _ __

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38

Homeostasis and Adaptation

39

Homeostasis and Adaptation

Tropisms

Plant Hormones
Like animals, plants use hormones to regulate their growth and
development. Plant hormones (phytohormones) are organ1c
compounds produced in one part of the plant and transported to
another part, where they produce a growth response. Hormones

are effective in very small amounts. There are five groups of

phytohormones: auxins (e.g. IAA), gibberellins, cytokinins,
ethene, and abscisic acid (ABA). Together they control the growth
of the plant at various stages of development.

Young Leaves and Buds Auxin (IAA) is

produced in the young leaves and buds.
IAA is a strong promoter of growth in stem
length and controls the differentiation of
tissues. It is the hormone responsible for
apical dominance: the growing leaves of
the apical bud synthesise IAA at
concentrations high enough to suppress
the growth of lateral buds below.

Shoot growth Cytokinins promote cell division.

They move from the roots to the leaves in the
transpiration stream and, although they do not
influence growth in the length of the stem, they
keep the shoot and root growth in balance.
Gibberellin promotes elongation in the region
just below the shoot tip (subapical region).
Fruit Ethene (ethylene) accumulates
in mature fruit to induce ripening.
Abscisic acid (ABA) is produced in ripe
fruit, inducing fruit fall. Cytokinins
made in the dividing cells of young
fruit are essential for growth.
Leaves Abscisic acid (ABA) is a
growth inhibitor made in the leaf
chloroplasts in response to water
stress. It acts on the guard cells,
causing stomatal closure and
thereby reduces water loss.

1-- Cytokinins move up

Old Leaves Ethene and abscisic acid

(ABA) are made in the old (senescent)
leaves. Ethene promotes leaf fall through
the development of a zone across the stem
where the leaf will break off (the abscission
zone). ABA promotes seed dormancy.
Although it reaches high concentrations in
senescent leaves, its exact role in leaf fall
is unclear- it appears to promote
abscission in only a few species. Together
with IAA, gibberellins (produced in the
chloroplasts, embryo, and young leaves),
delay the onset of senescence and leaf fall.

E en though most plants are firmly rooted in the ground, they are
c:pable of growth movements and responses to environmental
timuli (cues). Some of these responses are slow and gradual
~hile others may be rapid and quite spectacular. Tropisms are
lant growth responses to external stimuli, where the stimulus
~irection determines the direction of the growth response.

Geotropism Growth responses to the "-.:::,__~~~

1
earth's gravitational pull. Stems and
coleoptiles are negatively geotropic
-they grow away from the direction of
the earth's gravitational pull.

Tropisms may be positive or negative depending on whether the

plant moves towards or away from the stimulus. The main stimuli
that cause growth responses in plants are gravity, light,
chemicals, and touch (pressure). Tropisms are distinguished
from nastic responses by the directionality of the response. The
direction of a nasty is independent of the stimulus direction.

Phototropism Growth reponses to light,

particularly directional light. Coleoptiles, young
stems, and some leaves are positively
phototropic. The receptor for the phototropic
response is probably a pigment or pigments
in the light sensitive tissues. These are
thought to trigger the redistribution of plant
hormones (auxin and /or a growth inhibitor) in
the region of cell elongation.

Thigmotropism Growth reponses to touch

or a solid surface. Tendrils (modified leaves)
have a coiling response stimulated by touch
and are positively thigmotropic

Chemotropism A growth response to

a chemical stimulus. Pollen tubes grow
towards the chemical released by the
ovule and away from the air (they are
negatively aerotropic).

the plant to the

\ shoot and leaves

Cambium Activity Auxin and gibberellins

promote cell enlargement and
differentiation in the cambium, promoting
the formation of secondary vascular
tissues (secondary thickening).

Seeds Gibberellins are involved in

breaking the dormancy of seeds and buds,
and in mobilising food stores during seed
germination. In some plants cytokinins are
also involved in seed germination.

Roots In mature plants, cytokinins

are synthesised in the root tips and
travel to the shoots and leaves in the
transpiration stream.

Root tip Auxin is synthesised in the

meristematic tissues of the plant: especially the
root and shoot tips, but also in the young leaves,
flowers and fruits. From these areas it is
transported to areas of growth in the plant.

Synthetic analogues of IAA Since the discovery of its chemical

structure, many analogues of IAA have been produced for commercial
use. As with IAA, these analogues are transported around the plant
where they exert an effect on its growth and metabolism. IAA
analogues are applied as growth promoters in rooting powders, and
as inducers for fruit production . Some analogues (e.g. 2-4-5-T) even
act as growth inhibitors and are used as selective herbicides.

Hydrotropism Growth response to water.

Roots are mainly influenced by gravity but
will also grow towards water and are said
to be positively hydrotropic.

Roots are positively geotropic, and curve downward after

emerging through the seed coat. The gravity-sensing
mechanism is probably based on the presence of statoliths small clusters of unbound starch grains in cells. These change
position in response to gravity, triggering the response through
the action of hormones (auxin and ABA have been suggested).

1. (a) Briefly define the term tropism: _________________________________________ _____________

1. Describe one commercial application of a plant hormone (name the hormone in you r answer):
Hormone: _____________________________
Application: ------------------- -------- -- - - - - - - - -

(b) Distinguish between a tropism and a nastic response: - - - - - - - - - - -- - -- - - - - - - - - - - -

2 . Explain the adaptive value of the following tropisms:

2. Explain the role of auxin (IAA) in the following plant growth processes:
(a) Apical dominance: ------------------------------------------------------------------

~)Pos~vegeotrop~minroo~:

______ _ _ _ _ _ _ _ __ __ _ _ _ __ __ _ _ _ _ _ _ __ __

(b) Positive phototropism in coleoptiles: - - - -- -- - - - - - - - - - - - - -- - - - - - - -- -

(b) Stem growth:

(c) Positive th igmotropism in weak stemmed plants, such as v i n e s : - - - - - - - - - - - - - - - - - - - - -

3. Explain why pruning (removing the central leader) induces bushy growth in plants: _ _______________ _______

(d) Positive chemotropism in pollen grains:--- - -- - - - - - - - - -- - - - - - - - - - - - (e) Negative geotropism in shoots: _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ __ _ __

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40

Homeo!?ta!?i!? and Adaptation

Homeo!?ta!?i!? and Adaptation

Investigating Phototropism
Phototropism in plants was linked to a growth promoting
substance in the 1920s. A number of classic experiments,
investigating phototropic responses in severed coleoptiles, gave
evidence for the hypothesis that auxin was responsible for tropic
responses in stems. Auxins promote cell elongation. Stem
curvature in response to light can therefore result from the
differential distribution of auxin either side of a stem. However,
the mechanisms of hormone action in plants are still not well

Investigating Geotropism

understood. Auxins increase cell elongation only over a certai n

concentration range. At certain levels, auxins stop inducing
elongation and begin to inhibit it. There is some experimental
evidence that contradicts the original auxin hypothesis and the
early experiments have been criticised for oversimplifying the
real situation. Outlined below are some experiments that
investigate plant growth in response to light and the role of
hormone(s) in controlling it (see also: Plant Hormones).

1. Directional Light: A pot plant is exposed to direct sunlight

near a window and as it grows, the shoot tip turns in the
direction of the sun. If the plant was rotated, it adjusted by
growing towards the sun in the new direction.

41

Although the response of shoots and roots to gravity is well

known, the mechanism behind it is not at all well understood. The
importance of auxin as a plant growth regulator, as well as its
widespread occurrence in plants, led to it being proposed as the
primary regulator in the geotropic response. The basis of auxin's
proposed role in geotropism is outlined below. The mechanism is
appealing in its simplicity but, as noted below, has been widely

criticised, and there is not a great deal of evidence to support it.

Many of the early plant growth experiments (including those on
phototropism) involved the use of coleoptiles. Their use has been
criticised because the coleoptile (the sheath surrounding the
young shoot of grasses) is a specialised and short-lived structure
and is probably not representative of plant tissues generally.

The Role of Auxins in the Geotropic Response of Stems and Roots

Directional Sunlight

Shoot grows in the

direction of sunlight

(a) Name the hormone that regulates this growth response:

Gravity

Gravity

) _.. /-;

~~:~

r----:
....
~
B

(b) Give the full name of this growth response:

(.:::<;:::__
Growing _ _ _.J.-_.,
shoot
of plant

(c) State how the cells behave to cause this change in shoot
direction at:
Point A : - - - - - - - - - - - - - - - - - Point 8 : - - - -- - - -- - - - - - - - - -

Draw your cells here:

(d) State which side (A or B) would have the highest

concentration of hormone:

A horizontally placed root

(radicle) tip grows downwards
- this is positive geotropism.

Experiments on isolated shoot tips provide some evidence that geotropism (like
phototropism) is due to different growth rates of upper and lower sides of the
stem or root in response to the redistribution of auxin. In a horizontally placed
shoot tip (see diagram, right), more auxin accumulates on the lower side than
on the uppermost side. In stems, this causes elongation of the cells on the lower
surface and the stem tip turns up. The root grows down because root elongation
is inhibited by the high levels of auxin on the lower surface (see graph below).

A horizontally placed stem

tip grows upwards -this is
negative geotropism

Auxin moves to the lower side.

The cells on the lower side
elongate in response to auxin
and the stem turns upwards.

Agar block
33% auxin
Barrier ---iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiii~

150

(e) Draw a diagram of the cells as they appear across the

stem from point A to 8 (in the rectangle on the right).

..

:l

.s"'
3l

2. Light Excluded from Shoot Tip: With a tin foil cap placed
over the top of the shoot tip, light is prevented from reaching it.
When growing under these conditions, the direction of growth
does not change towards the light source, but grows straight
up. State what conclusion can you come to about the source
and activity of the hormone that controls the growth response:

In a horizontally placed seedling, auxin moves to the

lower side of the organ in both the stem and root.
Whereas the stem tip grows upwards, the root tip
responds by growing down. Root elongation is inhibited
by the same level of auxin that stimulates stem growth
(see graph left). The higher auxin levels on the lower
su rface cause growth inhibition there. The most
elongated cells are then on the upper surface and the
root turns down. This simple auxin explanation for the
geotropic response has been much criticised: the
concentrations of auxins measured in the upper and
lower surfaces of horizontal stems and roots are too
small to account for the growth movements obseNed.
Alternative explanations suggest that growth inhibitors
are also somehow involved in the geotropic response.

100

l~l

..

c:

Directional Sunlight

Auxin Concentration and Root Growth

The auxin concentrations

that enhance stem growth - - - 4
inhibit the growth of roots

.c:

E
0

:; a:

- T i n foil cap

e>
0

~ ~l
~ ~

e> .S

c:

jjj

1Q- 5

10--3

1Q- 1

103

101

Increasing concentration of auxin mg 1- 1 (log 10 scale)

Growing
shoot ---+--<~

of plant

1. Explain the mechanism proposed for the role of auxin in the geotropic response in:
3. Cutting into the Transport System: Two identical plants were
placed side-by-side and subjected to the same directional light
source. Razor blades were cut half-way into the stem , thereby
interfering with the transport system of the stem. Plant A had
the cut on the same side as the light source, while Plant 8 was
cut on the shaded side. Predict the growth responses of:
Plant A: _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __

(a) Shoots ( s t e m s ) : - - - - - - - - - - -- -- -- - - - - - - - -Directional Sunlight

~
.

(b) Roots: _ _ _ _ _ _ _ _ _ __ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

A
PlantS: _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ __
Growing shoot
of plant

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--------

Razor blade
left in cut

2. (a) From the graph above, state t he auxin concentration at which root growth becomes inhibited: _ _ _ _ _ _ _ __
(b) State the response of stem at this concentration: _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
3. Briefly state a reason why the geotropic response in stems or roots is important to the survival of a seedling:
(a) Stems: ______________ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _____ _ ___
(b) Roots: - - - - - - - - - - - -- - - -- - - - - - - - - - - - - - --

Biozone International 2000

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- - -- - - -