Plant defences against herbivory include a

range of mechanical and chemical obstacles

suchas unpalatableor indigestiblechemicals,
toxins that kill or repel herbivores and
thorns. Defences can be constantly present
(constitutive) or produced by the plant
following damage or stress (induced).
Plants have also evolved indirect defensive
features by evolving ways of attracting the
natural enemies of the herbivores that feed
on that specifc plant. For example, the plants
emit odours that attract thesenatural enemies,
and when they arrive, the plants provide food
Scienceis all about testinghypotheses, andin
the June 2014 issue of Veld & Flora there was
a very interesting letter by Ralph Peckover on
p. 92. Ralph had noticed that some of his aloes
had many more spines on their leaves than
others of the same species, and he wondered
if this was caused by a reaction to continuous
browsing on the leaves by kudus. He writes,
It is a well known fact that kudus browsing
shrubs and trees cause the shrubs to warn
their neighbours that their leaves are being
eaten so they can increase their leaf tannins to
make themless palatable so the kudus move
away. But perhaps one should question this
well knownfactbecausethereisnoconclusive
scientifc research that substantiates the claim
that plantstalk to each other using tannins.
Tannin is a name for a large group of
polyphenolic compounds that make up just
some of the macromolecules that are stored
inplant cells. Plants producetannins andother
longmoleculecompounds (many of whichare
insoluble andhence very stable insitu) that are
packed full of carbon, hydrogen and oxygen
(CHO) because they are cheap molecules to
make and enable the plants to dump excess
CHOmolecules. Oneof thepropertiesof tannin
is that it binds protein (and so limits protein
absorption in animals), hence our conclusion
that they function as anti-herbivore defences.
But is this correct? Recently ecologists have
found that tannins are important controllers
of decomposition and nitrogen recycling in
the soil, and this may well be their primary
function not defence, which is a spin-of. The
only way plants can rapidly absorb molecules
from the air is through their stomata those
tiny apertures that allow atmospheric gases
to enter and leave the plant. It is unlikely that
macromolecules, that are insoluble, are going
to be an efective means of communication.
l earni ng about bi odi versi ty Veld & Flora FACTSHEET PLANT DEFENCES
Plant defences
and shelter for them; as in the case of biting
ants associated with acacias. Herbivory may
also be encouraged by some plants to assist in
their reproduction notably the productionof
nectar to attract bees and other pollinators to
performessential cross-pollination.
Herbivory is not always bad for a plant
and in some cases it provides a positive
stimulus for plant growth. This is the case
for the majority of grass species that grow
much faster if they are constantly cropped by
grazers such as the huge wildebeest herds of
the Serengeti plains.
Chemical warfare
Mechanical defences
Can plants raise the alarm?
We know that plants in nutrient-poor
environments (such as our Cape fynbos)
have much higher tannin contents than
communities on richer soils. (Hence the tannin
stained waters of the Cape mountains and
the whisky making waters of Scotland.) Why
this occurs is that while the sun is shining and
plants haveaccess towater andcarbondioxide,
they are not able to switch of photosynthesis.
But protein building elements, Phosphorus(P),
Nitrogen (N) and Potassium (K), are limited
so plants in these low nutrient environments
have to fnd a way to dispose of the excess
CHO molecules from photosynthesis. Plants,
unlike animals, cannot excrete so they make
stable CHOmacromolecules and deposit them
in refuse areas. Co-incidentally, some of these
macromolecules also have anti-herbivore
properties. However, it is much more likely
that plants with macromolecules that aided
survival such as those molecules aiding
nitrogen re-cycling were selected through
evolution (hence the evergreen, sclerophyllous
vegetationof theCapemountainspackedfull of
tannins, oils andother CHOmacromolecules).
And how many of us have been told that
African thorn-trees have spines and thorns to
protect themfromherbivory whentheir major
herbivores areinsects?Thegirafeandelephants
I see eating the thorniest of plants seemto have
little difculty dealing with them. So could not
the thorns be just another CHOrefuse area? To
gobacktoRalphPeckoversletter, whereadjacent
aloes of the same species but from a diferent
regionwerenot browsed, this couldjust as easily
be that they are genetically slightly diferent
and so pre-programmed to produce more CHO
refuse locations (in this case, more spines). The
suggestion for further experimentation is great
however, the experimental plants must all be
fromthesamegenome.
WHAT DOES THAT MEAN?
Autotrophes Organisms capable of
synthesizing their own food from simple organic
substances such as higher plants, algae and
photosynthesizing bacteria.
CHOmolecules are the product of photosynthesis
usingthesuntocombinewater andcarbondioxide,
which are generally freely available i.e. not
limited like some other key elements for making
proteins like phosphorus, nitrogen and potassium.
Glucose, the frst CHO molecule synthesised in
photosynthesis, is the smallest building block for
all subsequent CHO molecules (some common
macromolecules are cellulose, lignin and tannin).
Hypothesis An idea or explanation that is tested
through study and experimentation.
Sclerophyllous Vegetation with small, evergreen,
tough, leathery leaves with thick cuticles.
TEXT and illustrations by Eugene Moll and
Caroline Voget, with input from The Story of Life
and the Environment: An African Perspective by Jo
van As, Johann du Preez, Leslie Brown and Nico
Smit (Struik Nature), and PlantZAfrica (http://
www.plantzafrica.com).
PHOTOS clockwise from top left: Acacia thorns
highly infated by ant activity. Photo: E. Moll. Girafes
browsingonUmbrellaThorn(Vachelliatortilis). Photo:
E. Moll. Sweet Thorn (Vachellia karroo) thorns. Photo:
E. Moll. Tree Euphorbia (Euphorbia triangularis)
not only has thorns and spines, but also exudes a
poisonous latex when the plant is damaged. Photo:
C. Voget. Buchu (Agathosma crenulata) leaf showing
oil glands. Photo: Alice Notten.
They key concept of evolution is that changes in populations are inheritable, and are
directed by natural selection. The process occurs within populations where individuals
gradually change in formor function to adapt to or to exploit newenvironmental conditions
or pressures. For example, this may include adaptive responses to predators, or defensive
adaptations to climate change.
In ecology, predation describes a biological interaction in which a predator organism
feeds on other living organisms known as prey. Animals that feed on living plants are called
herbivores. So herbivory is a formof predation in which a herbivore consumes autotrophes
(organisms capable of synthesizing their own food fromsimple organic substances) such as
higher plants, algae and photosynthesizing bacteria. Following that defnition, many fungi,
some bacteria andprotozoans anda small number of parasitic plants can also be considered
herbivores alongwithanimals. However, herbivorygenerallyapplies toanimals eatingplants.
Fungi, bacteriaandprotozoans that feedonlivingplants areusually termedplant pathogens.
Herbivores dependonplants for foodandhaveco-evolvedmechanisms toobtainthis food
despite the evolution of a diverse arsenal of plant defences against herbivory. Herbivores
adapt to plant defences to improve their chances of feeding on the plants. Plants, on the
other hand, protect themselves by limiting the ability of herbivores to eat them.
Plants have many external structural
defences that discourage herbivory. These
include structural defences on stems
and leaves like thorns. Some defensive
compounds are produced internally but
are released onto the plants surface; for
example, resins, thereby making feeding
difcult. Some plants produce sticky sap that
traps insects. Some are covered with sharp,
barbed hairs often containing irritants.
These all restrict the herbivores feeding
rate. It is thought that the whole structure
of a typical plant, its branching and leaf
arrangement, may also be evolved to reduce
herbivore impact. The African acacias are
well known for their thorns, but thorns and
spines may not be quite what we think they
are. See Prof. Molls text box on the right.
LINKS TO THE CURRICULUM
For Grade 12, Term 3, Strand 2: LIFE PROCESSES
IN PLANTS AND ANIMALS, section: Responding
to the environment: plants includes Plant defence
mechanisms: chemicals, thorns etc.
Plants fight back
Insects are a plants main enemy, and plants
have developed many chemical defences
against them. The Wild Peach (Kiggelaria
africana) has leaves that, when crushed,
smell like almonds the characteristic smell
of cyanide. And indeed, its leaves contain
cyanide, and only one butterfy larva, the
Garden Acraea, has managed to evolvea way
of safely eating them. It stores the cyanide
and is consequently not itself preyed upon,
except by some cuckoos that have evolved
ways of eating them and surviving the dose
of cyanide. Likewise, the almond-scented
leaves of the Wild Pear (Olinia cymosa)
also smell faintly of almonds and also
contain cyanide.
Buchus (Agasthosma) belong to the
Rutaceae, or citrus family, and are virtually
immune from attack by insects. This is due
to oil glands on the leaves which secrete
an aromatic smell. Agathosma betulina and
A. crenultata are commercially grown for
their oil which is extracted and used in the
manufacture of cosmetics and medicines.
121 SEPTEMBER 2014 | VELD&FLORA 120 VELD&FLORA | SEPTEMBER 2014