A258 Cereals C4 Plants

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B io Factsheet

www.curriculum-press.co.uk Number 258

Biology of cereals 1: Sorghum, a C4 plant


This Factsheet: These adaptations are reflected in the water use efficiency of
• Summarises the importance of cereals worldwide sorghum (Table 2)
• Describes and explains the adaptations of sorghum, a C4 plant Table 2. Water use efficiency of sorghum
• Reviews the very many types of exam questions that are set on
sorghum in particular and cereals in general Cereal Water use efficiency/ kg water used per kg
dry mass produced
Cereal crops such as sorghum are a major source of nutrients all Sorghum 300
over the world. Cereals grow well in many different environments,
their grains are easy to store, contain a wide variety of nutrients Wheat 350
and are often high in particularly important ones such as vitamin B. Maize 550
They also have high energy and fibre content.
Typical exam Question
Sorghum Scientists investigated the effect of the water-holding capacity
(WHC) of soil on the yield of sorghum and maize. Seeds of both
plants were planted in plots containing high WHC soil and in
plots containing low WHC soil. The yield of sorghum and maize
from all the plots was measured at the end of the investigation.
The graph shows the results.
6000

Tracey Slotta @ USDA-NRCS PLANTS Database 5000

Sorghum is a cereal that is well adapted to survive in hot, dry


yield/kg 4000
conditions (Table.1).
per hectare

Table.1 Adaptations of sorghum 3000

Feature Explanation
2000
Extensive, deep root system Maximises water absorption
Curled, small leaves Reduces water loss by 1000
evaporation/transpiration
Reduces surface area over 0
which transpiration can occur sorghum sorghum wheat wheat
LWC HWC LWC HWC
and traps layer of moist air
Waxy leaves composed of esters Wax doesn’t melt and shiny (a) Suggest three factors that would have to be controlled during
and fatty acids with a melting surface reflects heat this investigation to ensure that the results were valid (3).
point much higher than non- (b) (i) Describe the results shown (2)
tropical plants (ii) Suggest an explanation for the results (2)
yield for sorghum;
Leaves have rows of motor cells Traps layer of moist air,
yield for wheat is better in HWC soil / little difference in
along the midrib of the upper reducing the diffusion
wheat not well adapted to dry conditions;
surface of the leaf, enabling the gradient
(ii) sorghum tolerant of dry soils/ adapted to dry conditions;
leaves to roll up.
WHC had much greater effect on wheat yield;
Reduced stomatal numbers and Reduces water loss by WHC had little effect on sorghum yield;
stomata in pits evaporation/transpiration (b) (i) yield for sorghum greater than yield for wheat;
Reduces wilting Soil pH;
Increased sclerenchyma and
Wind/shelter;
silica
Aspect;
Tolerant of high temperatures and Can grow in tropical Watering regime;
heat shock proteins (HSPs) conditions HSPs help prevent Planting depth;
activate at very high temperatures damage to proteins Planting pattern/density;
(a) No. of seeds sown;
Markscheme
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258. Biology of cereals 1: Sorghum, a C4 plant Bio Factsheet
www.curriculum-press.co.uk

Photorespiration
Photorespiration isn’t on many specifications but that doesn’t stop the Examiners from setting questions on it – they often don’t mention
the term but give you a diagram of it that tests your ability to apply biological principles that are on the spec to a topic that isn’t. In other
words, they are testing your ability to think on your feet. Nice people these Examiners!
In the light-independent stage of photosynthesis, the enzyme ribulose bisphosphate carboxylase (Rubisco) catalyses the combination of
carbon dioxide with ribulose bisphosphate (RuBP).
The 6C compound formed immediately breaks down into two 3C compounds ( GP) which are then converted into triose phosphate ( (TP)
using ATP and NADPH from the light dependent stage (Fig. 1)

Fig. 1 The Calvin cycle (C3 pathway)

CO2

Light Dependent Stage


CO2 + RuBP occurs in thylakoids
Rubisco catalyst
carbon fixation 6C Unstable &
breaks down

2GP
(Glycerate-3-phosphate)

Regeneration of RuBP
ATP
(from Light NADPH
dependent stage) (from Light
dependent stage)

ADP
NADP

2TP
(Triose phosphate) Light Independent Stage
(Celvin Cycle) occurs in stroma
Synthesis of polysaccharides,
amino acids, lipids and nucleic
acids

However, Rubisco can also bind with oxygen. In hot, dry conditions plants close their stomata to prevent excess water loss. But this also
stops CO2 entering the leaf. When CO2 levels inside the leaf fall to low levels Rubisco combines with O2 instead of CO2. This process is
called photorespiration. It is called photorespiration because:

• It occurs in light
• It requires oxygen, like aerobic respiration
• It produces CO2 and water

However, it does not produce ATP and it reduces carbon dioxide assimilation because some of the intermediates in the Calvin cycle are
degraded. Photorespiration therefore reduces crop yields.

• GP is the first stable product of the light-independent stage. It is a three- carbon compound and for this reason the Calvin cycle is also
called the C3 pathway.

• Plants such as sorghum are C4 plants. They have evolved an additional way of initially fixing the carbon dioxide and preventing
photorespiration.

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258. Biology of cereals 1: Sorghum, a C4 plant Bio Factsheet
www.curriculum-press.co.uk

Fig. 2 The C4 pathway


CO2

1. Within the leaf mesophyll


cells, instead of CO2 being 1
fixed by the enzyme 2. Oxaloacetate is
reduced by NADPH
Rubisco and combined (3C) Phosphoenol Olalocetate (4C)
with ribulose pyruvate into malate.
bisphosphate, it is fixed
by the enzyme NADPH
Mesophyll cell

phosphoenol pyruvate AMP


carboxylase (PEPC) and 2
combined with NADP
ATP
phosphoenol pyruvate
(PEP) to form the four- 3. The malate then moves out
5
carbon compound of the mesophyll cells into
oxaloacetate (hence, C4 the bundle sheath cells that
Pyruvate (3C) (4C) Malate
pathway). are tightly packed around
the veins of the leaves.

5. The pyruvate re-enters 3


the mesophyll cells and
recats with ATP, Pyruvate (3C) (4C) Malate 4. Inside the bundle sheath cells a
regenerating PEP. decarboxylase enzyme converts
malate into pyruvate and
Bundle sheath cell

releases CO2. The levels of CO2


4 NADP
CO2 inside the bundle sheath cells
become extremely high: 10 – 60
NADPH
times the CO 2 concentration
Calvin found in mesophyll cells. This
Polysaccharides, then enables the normal Calvin
amino acids, Cycle
(C3) pathway to begin i.e.C4
lipids and plants exploit both a C4 and the
nucleic acids normal C3 pathway.

Vein

Fig. 3 Transverse section through the leaf of a C4 plant So what is the point of all this?
PEPC has an extremely high affinity for carbon dioxide, binding it
even when carbon dioxide levels are very low, and it has no affinity
epidermis/epidermal cell at all for oxygen. It also has a high optimum temperature – it
works best in hot conditions (45oC), precisely when the plant
may have closed its stomata, thus limiting the amount of CO2 that
can enter. In fact the enzyme is so efficient that C4 plants can
afford to keep their stomata closed (preventing water loss) much
longer than C3 plants can.
The tightly-packed mesophyll cells surrounding the bundle sheath
cells don’t let oxygen into them so Rubisco can’t bind to it when
CO2 levels are low.
Thus, C4 crops such as sorghum, maize and sugar cane are able
to withstand higher temperatures and light intensities, lose less
water via transpiration and hence provide greater yields than C3
crops
mesophyll (cell)
bundle sheath (cell) Extract from Chief Examiner’s Report:- Many candidates could
not explain why the C4 pathway enables plants to grow well in
hot conditions. Many candidates seemed to believe that the C4
pathway was a modified form of respiration.

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258. Biology of cereals 1: Sorghum, a C4 plant Bio Factsheet
www.curriculum-press.co.uk

Typical Exam Question (d) Species X has evolved an altered form of photosynthesis that
(a) The diagram shows sections through a typical leaf of three helps it to concentrate carbon dioxide in some cells. The diagram
different cereal plants. shows the interconversion and transfer of some organic
Cereal A Cereal B Cereal C compounds, which contain either three carbon atoms (C3) or
cuticle four carbon atoms (C4), in a leaf of species X.
CO2
Cell A Cell B

C3
compound C3
compound
AT P
stomata C4 C4 C4
CO2
compound compound compound
Which cereal is most likely to be grown in hot, dry conditions?
Explain your answer (2).
(b) The diagram shows a section of the lower leaf epidermis of Use information in the diagram to explain:
sorghum. (i) how reactions in cell A result in a large amount of carbon
dioxide to enter cell B (2)
(ii) how very high concentrations of carbon dioxide are built
up in cell B (2)
(iii) the significance of the fact that the stomata of species X
are only open for approximately 65% of the time that those
of species Y are open (3).

Maintaining high rate of photosynthesis;


Decarboxylation in cell B maintains high CO2 concentration;
But also stops CO2 diffusion into the leaf;
Stomatal closure reduces this;
(iii) High temperatures may result in high transpiration/ water loss;
Ref to use of ATP energy to transfer C3 compound into cell A;
accelerates decarboxylation/ ref to feedback control;
0.1 mm Movement of the C3 compound from B to A is rapid and
Sorghum has relatively few stomata. Explain the significance rapidly in B;
of this (2) (ii) Decarboxylation of the C4 compound is rapid/ CO2 is released
Allowing more CO2 to diffuse in;
(c) The graphs show the effects of light intensity and temperature Maintains the concentration gradient;
on the rate of photosynthesis in two crop species of plant, X low;
and Y. (i) C3 reaction rapidly fixes the CO2 / keeps CO2 concentration (d)
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30oC / temperature is still limiting at high temperature;
X Higher rate of photosynthesis at high temperature / Y slows above
5
Rate of is still limiting at high intensity;
photosynthesis 4 Higher rate of photosynthesis at high light intensity/ light intensity
/arbitary units 3 Species X; (c)
Y reduces transpiration/water loss; ref to C4;
2
grows in hot, dry /arid conditions; (b)
1 Reduce water loss;
0 Leaf A; Thick cuticle; Few stomata; Sunken stomata; (a)
0 10 20 30 40 50 60 70 Markscheme
Light intensity/arbitary units

6
5 X
4
3
Rate of
photosynthesis 2 Extract from Chief Examiner’s Report
/arbitary units Y (d) This part of the question was simply about diffusion but
1
many candidates appeared to “give up” apparently because
0 they were intimidated by the unfamiliar diagram. Most
0 10 20 30 40
Temperature /0C candidates did not even mention diffusion, nor the maintenance
of concentration gradients; candidates need more experience
One of the crops is well adapted to growing in tropical of applying simple biological principles to unfamiliar
conditions. Suggest, with reasons whether this is species X or material.
Y (2)

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258. Biology of cereals 1: Sorghum, a C4 plant Bio Factsheet
www.curriculum-press.co.uk

Scientists are collaborating worldwide to try to understand how (a) Explain how this leaf anatomy helps the plant achieve high
sorghum can tolerate the high temperatures that would kill many rates of carbon fixation at high temperatures (6).
other important crop plants. One suggestion is that their heat
tolerance is due to a substance known as GB. (b) Germinating seeds use the enzyme alpha amylase to break
down starch. Scientists investigated the effect of temperature
Scientists investigated this by growing two varieties of sorghum, on alpha amylase in seeds of sorghum and rice that were
one containing high levels of GB and one containing low levels of germinating. The graph shows the results.
GB, in both distilled water and in sodium chloride solution (sorghum
is often grown in irrigated water that contains sodium chloride). 4
They measured the rate of photosynthesis of all the plants. The
table shows the results.
3
Variety of sorghum Rate of photosynthesis / arbitrary units enzyme
Distilled water NaCl solution activity
/arbitrary 2
High GB 31 18 units
sorghum
Low GB 33 8 1
rice
What does this data tell us?
• The scientists wanted to find out whether GB levels affected 0
10 20 30 40 50 60 70 80
photosynthesis, hence yield temperature / 0C
• So they selected varieties with high and low GB levels and then
grew them in distilled water (no NaCl ) and in water that was (i) Describe the results (3)
similar to the irrigation water (with NaCl) (ii) What do these result suggest about the tertiary structure
• Simple Biology tells us that, normally, plant roots growing in of alpha amylase in sorghum and rice? (3)
salty water will have water pulled out of them by osmosis. Worse,
so alpha amylase in sorghum may have less H bonds;
in hot conditions, irrigation water will evaporate, making the high temperatures affect H bonds more than other types of bond;
remaining soil even saltier able to form more enzyme-substrate complexes;
• In the NaCl solution the variety with high GB achieved more stable;
than twice the rate of photosynthesis of the variety containing (ii) tertiary structure/active site of alpha amylase of sorghum more
low GB alpha amylase in sorghum has higher maximum activity;
• This suggests that the GB is lowering the water potential of the alpha amylase in sorghum has higher optimum temperature;
sorghum roots, stopping water being withdrawn osmotically, alpha amylase in sorghum has higher activity at all temperatures;
enabling the plant to absorb water, survive and grow (b) (i) in both, enzyme activity increases and then falls;
photorespiration also destroys compounds in the Calvin cycle
Heat shock proteins from which biomass is produced;
Despite sorghum being able to tolerate and take advantage of high photorespiration prevents carbon dioxide entering the Calvin cycle;
temperatures, yields of sorghum can be severely reduced if it is photorespiration is avoided;
exposed to very high temperatures during germination or at the PEP carboxylase not denatured;
young seedling stage. / no affinity for O2;
PEP carboxylase has high optimum temperature / greater affinity for
When sorghum seedlings are exposed to temperatures above 37°C, sheath cells;
they produce heat shock proteins ( HSPs ). These intracellular decarboxylation of malate maintains high CO2 concentration in bundle
proteins help protect enzymes against denaturation and help prevent malate moves from mesophyll cell to bundle sheath cell;
damage to proteins as they are being synthesised. However, very light independent stage/Calvin cycle occur in bundle sheath cells;
young sorghum seedlings are unable to activate sufficient HSPs prevent O2 cannot reaching bundle sheath cells;
and this is why they are particularly vulnerable. 2. (a) tightly packed mesophyll cells;
Reduces area from which water can evaporate;
Practice Questions Rolling of leaves;
1. Outline how sorghum is adapted to growing in hot, dry Allows water to be collected from large area;
conditions (6) Dense/wide/deep root system;
temperatures;
2. The diagram shows a transverse section through the leaf of a
Allows photosynthesis when stomata closed/more efficient at high
C4 plant.
epidermis/epidermal cell C4 pathway;
Adult/embryo plants able to tolerate high temperatures;
Reduces water loss;
Sunken stomata;
1. Thick waxy cuticle;
Markschemes
Acknowledgements:
This Factsheet was researched and written by Kevin Byrne.
Curriculum Press, Bank House, 105 King Street, Wellington, Shropshire, TF1 1NU.
Bio Factsheets may be copied free of charge by teaching staff or students, provided that their
school is a registered subscriber. No part of these Factsheets may be reproduced, stored in
a retrieval system, or transmitted, in any other form or by any other means, without the prior
bundle sheath (cell) mesophyll (cell) permission of the publisher. ISSN 1351-5136
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