6

CHAPTER
Plant Adaptations
to the Environment
© 2016 Pearson Education, Ltd.

Elements of ECOLOGY Lecture Presentation by

NINTH EDITION, GLOBAL EDITION
Carla Ann Hass
Penn State University
Thomas M. Smith • Robert Leo Smith
 Chapter 6 Plant Adaptations to the
Environment
§ All life on Earth is carbon-based.
§ What does this statement mean?
§ What is the source of carbon from which life is
constructed?

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ What is the process of photosynthesis?
§ How is carbon dioxide in the atmosphere converted
to the mass of a plant?

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ Photosynthesis – light energy from the Sun drives a
series of reactions that result in the fixation of CO2
and the release of O2
§ Which wavelengths of light are used in this process?

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ Photosynthetic pigments absorb light in the
400–700 nm range, about the same as the visible
spectrum for humans
§ photosynthetically active radiation (PAR)
§ This light energy is converted to the chemical bond
energy in sugar:
6CO2 + 12H2O → C6H12O6 + 6O2 + 6H2O
§ Other carbon-based molecules including complex
carbohydrates, proteins, and fatty acids are made in
other parts of the plant

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ How is light energy converted to chemical bond
energy?
§ What types of chemical reactions are involved?

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ The light-independent reactions convert light energy
into chemical bond energy
§ Chlorophyll is a pigment in the chloroplasts that
absorbs light energy
§ Energy is transferred to an acceptor
§ Photosynthetic electron transport:
§ synthesizes ATP
§ reduces NADP+ to NADPH
§ These two molecules carry energy to the light-
independent reactions

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ The light-dependent reactions use the chemical
bond energy in ATP and NAPDH to incorporate CO2
into simple sugars
§ They do not directly require sunlight to take place
§ How is a molecule that is an atmospheric gas used
to produce a solid molecule?

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ During carboxylation:
§ RuBP (a five-carbon molecule) combines with CO2
and fixes it to a solid form
§ this reaction is catalyzed by an enzyme, rubisco
§ The reaction produces two molecules of 3-PGA (a
three-carbon molecule): C3 cycle
§ 3-PGA, ATP, and NADP are used to synthesize the
energy rich sugar G3P
§ What is the fate of G3P?

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ The fate of G3P
§ some is used to produce sugars and other
carbohydrates
§ glucose
§ starch
§ most is used to synthesize new RuBP
§ requires ATP
§ Light availability can limit the light-independent
reactions because it controls ATP and NADPH
production

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ Summary of C3 Photosynthesis

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Figure 6.1

Mesophyll cells
Chloroplasts

Stoma
Vascular
bundles

CO2 Light
(PAR)

CO2
Rubisco

From light RuBP

3-PGA
reactions
Calvin ATP Light
ATP cycle NADPH reactions

G3P

Simple sugars
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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ Plants use cellular respiration to convert the energy
in sugars and other molecules into ATP
C6H12O6 + 6O2 → 6CO2 + 6H2O + ATP
§ They are both using CO2 during photosynthesis and
producing it during respiration
§ Net photosynthesis = Photosynthesis - Respiration
§ Usually measured in moles CO2 per leaf area (or
mass) per unit time
§ µmol/m2/s

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 Section 6.1 Photosynthesis Is the Conversion
of Carbon Dioxide into Simple Sugars
§ The C3 pathway has one major drawback
§ Rubisco is also an oxygenase
§ catalyzes the reaction between O2 and RuBP
§ results in photorespiration – the release of CO2
§ Reduces the efficiency of C3 photosynthesis by as
much as 25%
§ Plants in high-light environments carry out other
types of photosynthesis that circumvent this problem

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 Section 6.2 The Light a Plant Receives
Affects Its Photosynthetic Activity
§ The availability of light directly influences the rate of
photosynthesis
§ What happens when it is dark?
§ Does the rate of photosynthesis continue to increase
as the level of light increases?

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 Section 6.2 The Light a Plant Receives
Affects Its Photosynthetic Activity
§ When it is dark and there is no PAR available
(= zero)
§ only respiration occurs
§ CO2 is lost (net uptake is negative)
§ Can use this point to estimate the rate of respiration
occurring in the leaf

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 Section 6.2 The Light a Plant Receives
Affects Its Photosynthetic Activity
§ As PAR increases, the rate of photosynthesis
increases
§ Light compensation point (LCP) is the light level
(PAR) at which
§ CO2 uptake in photosynthesis = CO2 loss in
respiration
§ Rate of net photosynthesis is zero

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 Section 6.2 The Light a Plant Receives
Affects Its Photosynthetic Activity
§ As light levels exceed the LCP, the rate of net
photosynthesis increases with PAR
§ Does this continue indefinitely?

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 Section 6.2 The Light a Plant Receives
Affects Its Photosynthetic Activity
§ Light saturation point = value of PAR above which
photosynthesis does not increase
§ Photoinhibition – the rate of photosynthesis will
decline as PAR exceeds the saturation point (seen in
some shade-adapted plants)

© 2016 Pearson Education, Ltd.

Figure 6.2

Maximum rate
of net photosynthesis
20
Net CO2 uptake (µmol/CO2/m2/s)

10 Light
saturation
point
Light compensation
point

0
500 1000 1500 2000
PAR (µmol/m2/s)

Rate of respiration
-10
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 Section 6.3 Photosynthesis Involves
Exchanges between the Plant and Atmosphere
§ The reactions of photosynthesis take place with
specialized leaf cells called mesophyll cells
§ What molecule in the atmosphere must be available
to these cells for these reactions to occur?
§ What molecule produced by these reactions may be
released to the atmosphere?

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 Section 6.3 Photosynthesis Involves
Exchanges between the Plant and Atmosphere
§ CO2 must move from the atmosphere into the leaf
§ Stomata – openings on the leaf surface that allow
CO2 to enter
§ gas diffuses from higher concentration in the air to
lower concentration in the leaf
§ at equilibrium, concentration is equal
§ Stomata are found in terrestrial plants
§ How is diffusion controlled?

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 Section 6.3 Photosynthesis Involves
Exchanges between the Plant and Atmosphere
§ Leaf cross section showing location of stomata

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Figure 6.3

Stoma
H2O CO2

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 Section 6.3 Photosynthesis Involves
Exchanges between the Plant and Atmosphere
§ Diffusion of CO2 is controlled by
§ the diffusion gradient = difference in CO2
concentration in the air next to the leaf and the leaf
interior (ppm)
§ stomatal conduction = the flow rate of CO2 through
the stomata (µmol/m2/s)
§ this is determined by
§ stomatal density – number per unit leaf surface area
§ aperture – size of stomatal openings

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 Section 6.3 Photosynthesis Involves
Exchanges between the Plant and Atmosphere
§ The stomatal aperture is under plant control
§ Stomata are open when
§ the concentration of CO2 in the air outside the leaf is
greater than that inside the leaf
§ CO2 in the leaf is constantly being converted into sugar
so the gradient is maintained
§ Stomata are usually closed when
§ photosynthesis and the demand for CO2 are reduced
for any reason
§ Why do the stomata close?

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 Section 6.3 Photosynthesis Involves
Exchanges between the Plant and Atmosphere
§ Transpiration – when the stomata are open, water
vapor in the leaf diffuses out
§ Transpiration rate depends on the diffusion gradient
and stomatal conduction of H2O vapor
§ the relative humidity inside a leaf is usually greater
than 99 percent
§ the lower the relative humidity of the air outside the
leaf, the larger the diffusion gradient and the more
rapidly the water will diffuse out of the leaf
§ How is this water replaced?

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Turgor pressure is the force exerted outward on a
cell wall by the water inside the cell
§ plants function best when their cells are fully hydrated
(at maximum turgor)
§ What happens to the plant when the water content
of its cells declines?
§ How do plants replace water lost through
transpiration?

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Plants take up water from the soil by their roots and
transport that water to their leaves
§ This occurs through passive transport so does not
require energy input but does require the transfer of
energy
§ Review from physics –
§ Gibbs energy (G) = energy available to do work
§ movement of water from soil to plant to atmosphere is
driven by internal differences in the G of water – a
gradient of decreasing energy

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Water potential (y) – describes the Gibbs energy of
water at any point in the soil-plant-atmosphere
continuum
§ Water potential = the difference in Gibbs energy per
mole between the water in the continuum and pure
water
§ It is expressed in terms of pressure (energy per
volume) using Pascals
§ Pa = 1 Newton/m2
§ y = zero for pure water at atmospheric pressure

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Examine the movement of water as a function of the
water potential gradient
§ Energy transfers always proceed from a region of
higher energy content to a region of lower energy
content
§ Start with the exchange of water between the leaf
and atmosphere
§ From the previous discussion of stomata, in which
direction does water vapor move?

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ When relative humidity = 100 percent:
yatm = 0
§ When relative humidity < 100 percent:
yatm becomes negative
§ As long as the relative humidity of the air is less than
99 percent:
yleaf > yatm

§ water moves from the interior of the leaf to the

atmosphere

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Atmospheric water potential is determined only by
humidity
§ Water potential within the plant is determined by
several factors
§ yp = hydrostatic pressure
§ This factor is the result of turgor pressure
§ turgor pressure increases a plant’s water potential
§ a decrease in turgor pressure due to water loss
decreases water potential

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ yp = osmotic potential
§ This factor is the result of solute concentration within
cells
§ The difference in solute concentration inside and
outside of the cell causes movement of water
through osmosis
§ increasing concentrations of solutes in plant cells
occur as a result of water loss and decrease water
potential

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ ym = matric potential
§ This factor is the result of large molecules, such as
those in the cell water, exerting an attractive force
on water
§ the tendency for water to adhere to a surface
(adhesion) decreases the water potential

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Total water potential at any plant, from the leaf to
the root is the sum of these:
y = yp + yp + ym
§ Osmotic and matric potentials are always negative
§ Turgor pressure may be positive or negative
§ Total potential can be either positive or negative but
values are typically negative

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Movement of water in a plant goes from areas of
higher water potential to lower potential
§ Depends on increasingly negative water potential as
water moves from soil to root to leaf to atmosphere:
yatm < yleaf < yroot < ysoil
§ There is daily variation in yleaf and yroot
§ stomata close at night and yleaf is less negative
because water is not being lost

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Figure 6.4
200

Atmospheric water potential (MPa)

For the continued movement of water from
the soil, into the roots, through the plant to
the leaves, and from the leaves to the 160 y atm
atmosphere through the process of
transpiration, the following condition must
hold: 120
y atm < y leaf < y root < y soil
80

Atmosphere 40

Transpiration 0
y atm 0 20 40 60 80 100
(a) Relative humidity (%)

0
y leaf
y soil
y root

Water potential (MPa)

-0.5
Transport of
water to leaf y leaf

-1.0

-1.5
Soil water
taken up
by roots y root -3.0
1 2 3 4 5 6
(b) Time (days)

y soil As the soil dries, y soil declines, requiring a

corresponding decline in y root and y leaf. As
all three values of water potential decline,
the gradient of increasingly negative water
potential from the soil, to the roots, and from
the roots to the leaves is maintained.

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ Water loss through transpiration will continue as
long as
§ light energy striking the leaf supplies enough heat for
evaporation
§ moisture is available in the soil
§ the roots are able to maintain a more negative water
potential than the soil
§ What affects the water potential of the soil?

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ At field capacity
§ water is freely available
§ ysoil is at or close to zero
§ As water is taken from the soil
§ ysoil becomes more negative
§ water holds more tightly to the soil
§ ym becomes more negative
§ This is strongly affected by soil type
§ clays soils have a higher surface area so have more
negativeym than sandy soils

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ If ysoil becomes more negative, how does this affect
yroot and yleaf?

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ yroot and yleaf must also become more negative to
maintain the water potential gradient
§ What happens if there is no rain so the soil water
continues to decrease? How does the plant reduce
water loss?

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Figure 6.4b

Atmosphere

Transpiration

0
y leaf
y soil
y root

Water potential (MPa)

-0.5
Transport of
water to leaf y leaf

-1.0

-1.5
Soil water
taken up
by roots
y root -3.0
1 2 3 4 5 6
(b) Time (days)

y soil
As the soil dries, y soil declines, requiring a
corresponding decline in y root and y leaf. As
all three values of water potential decline,
the gradient of increasingly negative water
potential from the soil, to the roots, and from
the roots to the leaves is maintained.

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ If there is no rain, eventually the gradient will be lost
§ At that point, the stomata close to prevent further
water loss through transpiration
§ What else is affected by the closing of the stomata?

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ If the stomata close, CO2 uptake stops
§ The value of yleaf at which net photosynthesis stops
varies between plants

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Figure 6.5

Net photosynthesis (mg CO2/dm2/h) 12

10 Speckled alder
(Alnus rugosa)

8
White ash
(Fraxinus americana)
6
Red oak
4 (Quercus rubra)

0
-1 -2 -3
y leaf, Leaf water potential (MPa)
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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ The rate of transpiration varies daily depending on
environmental conditions and the characteristics of
an individual plant
§ Controlling the stomata is a plant’s most important
way to regulate water loss
§ leads to a trade-off between taking in CO2 for
photosynthesis and losing water that it requires to live
§ water use efficiency = the ratio of carbon fixed per
unit of water lost

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 Section 6.4 Water Moves from the Soil,
through the Plant, to the Atmosphere
§ The balance between photosynthesis and
transpiration is an important evolutionary constraint
in terrestrial plants
§ Directly influences the productivity of different
ecosystems with different environmental conditions

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 Section 6.5 The Process of Carbon Uptake
Differs for Aquatic and Terrestrial Autotrophs
§ Aquatic autotrophs include submerged plants, algae,
and phytoplankton
§ Do aquatic autotrophs have stomata?
§ How do they acquire the CO2 needed for
photosynthesis?

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 Section 6.5 The Process of Carbon Uptake
Differs for Aquatic and Terrestrial Autotrophs
§ Recall:
§ CO2 diffuses from the atmosphere into surface
waters and mixes into the water column
§ Carbon dioxide – carbonic acid – bicarbonate
system tends to stay in equilibrium
CO2 + H2O H2CO3
H2CO3 HCO-3 + H+
HCO3- H+ + CO32-

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 Section 6.5 The Process of Carbon Uptake
Differs for Aquatic and Terrestrial Autotrophs
§ In aquatic autotrophs, CO2 diffuses from the water
across the cell membrane into the cell
§ Some aquatic autotrophs can use bicarbonate as a
carbon source
§ convert HCO3- to CO2 using the enzyme carbonic
anhydrase either by
§ active transport of bicarbonate into the cell, then
conversion
§ excretion of the enzyme into the surrounding waters,
then uptake of converted CO2

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Both photosynthesis and cellular respiration respond

to temperature variation, increasing as temperature
increases
§ What determines the maximum rate relative to
temperature for photosynthesis?

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ The temperature response of rubisco determines the

maximum rate of photosynthesis
§ The maximum temperature for photosynthesis is
lower than the maximum temperature for cellular
respiration
§ The overall temperature response of net
photosynthesis = rate of carbon uptake
(photosynthesis) - rate of carbon loss (respiration)

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ The temperature response curve is described by

three values:
§ Tmin = minimum temperature
§ (net photosynthesis near zero)
§ Tmax = maximum temperature
§ (net photosynthesis near zero)
§ Topt = range of temperatures over which net carbon
uptake is highest

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Figure 6.6

Net photosynthesis
Topt

(µmol/m2/s)
Tmin
Tmax
-10 0 25 45
Temperature (°C)
CO2 (µmol/m2/s)

Gross Respiration
photosynthesis

-10 0 10 20 30 40 50 60
Temperature (°C)
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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Leaf temperature determines the rates of these two

processes, not the air temperature
§ Plants
§ absorb both shortwave (solar) and longwave
(thermal) radiation
§ reflect some solar radiation and emit some longwave
back to the atmosphere
§ Net radiation balance
§ Rn = radiation received - radiation reflected/emitted to
environment

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Usually less than 5 percent of Rn is used for

metabolic processes and stored in chemical bonds
§ Remainder raises the temperature of the leaves and
surrounding air
§ What happens to the temperature inside a leaf on a
clear and sunny day?

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ On a clear and sunny day, internal leaf temperature

can be much higher than air temperature and may
reach critical levels
§ To maintain conditions for positive net
photosynthesis plants must exchange heat with the
environment
§ Second Law of Thermodynamics – energy flows from
areas of higher temperature to areas of lower
temperature
§ How does this heat loss occur in plants?

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Terrestrial plants lose heat by evaporation and

convection
§ Evaporation – part of transpiration
§ it requires a large amount of energy to change water
from a liquid to a gas
§ water transpires from the leaves and thermal energy
is lost
§ What is the relationship between the rate of
transpiration and the rate of evaporative cooling?

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Terrestrial plants lose heat by evaporation and

convection
§ Evaporation – part of transpiration
§ it requires a large amount of energy to change water
from a liquid to a gas
§ water transpires from the leaves and thermal energy
is lost
§ The higher the rate of transpiration, the greater the
evaporative cooling

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Convection is the transfer of heat energy through

the circulation of fluids (air/water)
§ Conduction is the transfer of heat energy from one
object to another by direct contact
§ If the surface of the leaf is warmer
§ it transfers thermal energy to adjacent molecules of
air or water through conduction
§ thermal energy then moves from the air next to the
leaf to the surrounding air through convection
§ Aquatic plants use convection to lose heat

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Boundary layer – the layer of still air or water next

to the surface of each leaf
§ It is different than the surrounding environment
because of the heat, water vapor and CO2 diffusing
from the leaf surface
§ if there is no air or water flow, the boundary layer
increases in thickness
§ this reduces the transfer of heat, water and CO2
between the leaf and the environment

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Leaf size and shape influence the boundary layer

thickness and dynamics
§ Air flows more smoothly over a larger continuous
surface
§ The boundary layer is reduced for
§ smaller leaves
§ deeply lobed leaves
§ compound leaves

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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ Sycamore leaves are larger and less lobed than oak

leaves so have a larger boundary layer and reduced
heat transfer

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Figure 6.7

oak

sycamore

(a)

39

36
Temperature (°C)

33 Oak
Sycamore

30

24
-90 -60 -30 0 30 60
Distance from center (mm)
(b)
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 Section 6.6 Plant Temperatures Reflect Their Energy
Balance with the Surrounding Environment

§ The physical environment also has an effect

§ Areas of high water availability
§ most heat is lost through transpiration
§ Areas of lower water availability
§ transpiration is limited to reduce water loss
§ most heat is lost through convection
§ plants tend to have smaller leaves
§ This illustrates another adaptive trade-off

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 Section 6.7 Constraints Imposed by the Physical
Environment Have Resulted in a Wide Array of Plant
Adaptations
§ There are many features of the physical
environment that influence plant processes
§ many are interdependent
§ What are some of these features and how do they
interact?
§ What types of plant adaptations are seen in different
environments?

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 Section 6.7 Constraints Imposed by the Physical
Environment Have Resulted in a Wide Array of Plant
Adaptations
§ Solar radiation influences
§ the amount of PAR
§ the temperature of the leaf and surrounding air
§ the temperature of the surrounding air affects the
relative humidity
§ relative humidity affects the rates of transpiration and
evaporation of water in the soil
§ What type of adaptations would you expect in plants
that live in dry, sunny environments?

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 Section 6.7 Constraints Imposed by the Physical
Environment Have Resulted in a Wide Array of Plant
Adaptations
§ A sunny, dry environment has
§ greater PAR
§ higher air temperatures
§ higher leaf temperatures
§ lower humidity
§ higher rate of transpiration
§ higher rate of evaporation
§ Plants adapted to this environment might have
smaller leaves, more leaves, more roots, deeper
roots

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 Section 6.7 Constraints Imposed by the Physical
Environment Have Resulted in a Wide Array of Plant
Adaptations
§ Multiple environmental conditions can lead to
adaptive trade-offs
§ adaptations for one environment may not be as
effective in a different environment
§ Plants distribute carbon acquired through
photosynthesis
§ allocating more carbon to producing leaves and
stems increases access to light and CO2
§ reduces the carbon available to produce roots,
decreases access to water and soil nutrients

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 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ The amount of solar radiation reaching the surface
of the Earth varies daily, seasonally and
geographically.
§ What is a major factor affecting the PAR that a plant
receives?
§ What type of adaptations to sun and shade are seen
in plants?

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 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ Shade intolerant plants – adapted to high-light
environments
§ Shade tolerant plants – adapted to low-light
environments
§ Differ across a wide variety of physical and
physiological characteristics that are adaptations to
sun and shade environments
§ Show fundamental differences in patterns of
photosynthesis in response to different levels of
available light

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 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ Shade environments – low light limits the rate of
photosynthesis
§ shade-adapted species produce less rubisco
§ require less energy so leaf respiration lower
§ light saturation point is lower
§ Moving a shade-tolerant plant to a high light
environment
§ its maximum rate of photosynthesis is limited
§ it will gain less carbon and grow less than a shade-
intolerant species

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Figure 6.8

20 Shade

Net CO2 uptake (µmol CO2 m2/s)

intolerant

Shade
10
tolerant
Light saturation
points

0
250 750 1250
PAR (µmol/m2/s)
Light
compensation
-10 points

(a)

16
Net photosynthesis (µmol CO2/m2/s)

14
Pinus nigra
12
10
8
6
Acer opalus
4
2
0
-2
-4

0 100 200 400 600 800 1000 1200 1400 1600 1800
PAR (µmol/m2/s)
(b)
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 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ Reich and colleagues examined nine species of
trees from cool temperate forests in North America
that vary in shade tolerance
§ Seedlings of all species were grown under the same
greenhouse conditions
§ Shade-intolerant species had lower levels of net
photosynthesis, leaf respiration, relative growth rate

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Figure 6.9

Maximum net photosynthesis

500

400

(nmol CO2/g/s)
300

200

100

0
1 2 3 4 5 6 7 8 9
(a) Species

35
30

Leaf respiration
25

(nmol/g/s)
20
15
10
5
0
1 2 3 4 5 6 7 8 9
(b)
Species

200
Relative growth rate

150
(mg/g/day)

100

50

0
1 2 3 4 5 6 7 8 9
(c)
Species

Shade tolerance
Low High
(shade intolerant) (shade tolerant)
© 2016 Pearson Education, Ltd.
 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ Augspurger looked at tree species from the tropical
rain forest of Panama
§ Grew seedlings of each species under
§ low light – shaded environment under forest canopy
§ high light – sunny environment under gaps in the
forest canopy cause by trees dying
§ Shade-intolerant species – survival and growth
greatly reduced in shade conditions
§ Shade-tolerant species – little difference in survival
and growth under the two conditions

© 2016 Pearson Education, Ltd.

Figure 6.10

Sun
Shade
20 1.5
Myroxylon balsamum

Seedling survival (log [number surviving +1])

10 0.9
Mean seedling height (cm)

0 20 40 0.3 Myroxylon balsamum

50 0 20 40
Ceiba pentandra Time (weeks)
40

30 1.5

20 0.9
10
0.3 Ceiba pentandra
0
0 20 40 0 20 40
Time (weeks) Time (weeks)
© 2016 Pearson Education, Ltd.
 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ Shade-tolerant and shade-intolerant species also
show differences in leaf morphology
§ Specific Leaf Area (SLA)
§ ratio of surface area in cm2 to weight in g
§ the surface area of leaf produced per gram of
biomass allocated to leaf production
§ In general, shade-tolerant plants have greater SLA.
Why?

© 2016 Pearson Education, Ltd.

 Section 6.8 Species of Plants Are Adapted
to Different Light Environments
§ Greater SLA increases the surface area for light
capture/unit of biomass
§ Leaf thickness may decrease as SLA increases
§ Plants also show phenotypic plasticity of SLA in
response to light levels in the environment
§ open – full sun
§ under canopy – shaded

© 2016 Pearson Education, Ltd.

 Based on these experimental results measuring SLA
and leaf thickness for shade-intolerant and shade-
tolerant plants both in the open and under the canopy,
which of the following is true?

A. both types of plants have higher SLA in the open.

B. shade-tolerant plants have lower SLA in both
places.
C. both types of plants have higher SLA under
canopy.
D. shade-intolerant plants have higher SLA in both
places.
E. SLA shows phenotypic plasticity but thickness
doesn’t.
© 2016 Pearson Education, Ltd.
Figure 6.11

500
450 Leaf thickness (µm)
400 SLA (cm2/g)
350
300
250
200
150
100
50
0
open under open under
Shade-intolerant Shade-tolerant
average average
© 2016 Pearson Education, Ltd.
 Based on these experimental results measuring SLA
and leaf thickness for shade-intolerant and shade-
tolerant plants both in the open and under the canopy,
which of the following is true?

A. both types of plants have higher SLA in the open.

B. shade-tolerant plants have lower SLA in both
places.
C. both types of plants have higher SLA under
canopy.
D. shade-intolerant plants have higher SLA in both
places.
E. SLA shows phenotypic plasticity but thickness
doesn’t.
© 2016 Pearson Education, Ltd.
 Quantifying Ecology 6.1 Relative Growth
Rate
§ How do you compare the growth rate of organisms
that are different sizes, or track the growth of one
individual through time?
§ Relative growth rate (RGR) – expresses growth as
a function of the size of an individual organism

© 2016 Pearson Education, Ltd.

 Quantifying Ecology 6.1 Relative Growth
Rate
§ Growth during a time period
§ grams (g) weight gained
§ Size of the individual at the beginning of the time
period
§ grams (g) weight at the beginning
§ To get a rate:
§ Grams weight gain/grams weight at beginning/time =
(g/g/time) = RGR

© 2016 Pearson Education, Ltd.

 Quantifying Ecology 6.1 Relative Growth
Rate
§ For plants, you can partition RGR into the different
components that contribute to growth
§ Net assimilation rate (NAR)
§ assimilation of new tissues per unit leaf area
(g/cm2/time)
§ function of total gross photosynthesis minus total
respiration expressed on a leaf area basis
§ Leaf area ratio (LAR)
§ leaf area per unit of plant weight (cm2/g)
§ the amount of assimilation allocated to make leaves
§ RGR (g/g/time) = NAR (g/cm2/time) ´ LAR (cm2/g)
© 2016 Pearson Education, Ltd.
 Quantifying Ecology 6.1 Relative Growth
Rate
§ Leaf area ratio (LAR) can be split into two parts
describing the allocation of net assimilation to leaves
§ Leaf weight ratio (LWR)
§ total weight of leaves/total plant weight
§ g leaves/g total plant weight
§ Specific leaf area (SLA)
§ measures leaf density or thickness
§ area of leaf/g leaf weight
§ if density is the same, a thinner leaf has greater SLA

© 2016 Pearson Education, Ltd.

 Quantifying Ecology 6.1 Relative Growth
Rate
§ Partitioning RGR allows for comparison between
different species or between individuals within the
same species when grown under different conditions

© 2016 Pearson Education, Ltd.

Quantifying Ecology Box Table 1

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ The demand for water is related to temperature
§ as air temperature rises
§ saturation vapor pressure rises
§ the rate of transpiration rises
§ What happens to the amount of water the plant
needs?
§ On what timescales does water availability vary?

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Plants open/close stomata depending on current
conditions
§ Stomata may be closed or partially closed
§ when the atmosphere is dry
§ when soil is dry
§ during the hottest part of the day
§ As water stress continues
§ stomata may be open only during cooler, more humid
conditions such as early morning
§ What happens when the stomata are closed?

© 2016 Pearson Education, Ltd.

Figure 6.12

35 100
Leaf
temperature

Relative humidity (%)

Leaf temperature (°C)
30
80

25

Relative 60
20 humidity

15 40
(a)

500
Stomatal conductance (mmol/m2/s)

400

300

200

100

0
6 8 10 12 14 16 18
Time of day (hour)
(b)
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Closing the stomata
§ reduces water loss through transpiration
§ reduces CO2 diffusion into the leaf
§ reduces evaporative cooling
§ Photosynthesis declines, leaf temperature can rise
§ Some plants respond to moisture stress by
§ curling leaves or wilting
§ reduces water loss
§ reduces heat gain because the surface area of the leaf
is reduced

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ What happens if water stress is prolonged?
§ chlorophyll production is inhibited and leaves turn
yellow
§ deciduous trees prematurely shed leaves
§ All of these changes represent acclimation to the
current conditions
§ How can water availability affect plant development?

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ If a plant encounters low soil water availability during
development, carbon allocation is
§ increased for the production of roots
§ decreased for the production of leaves
§ How does this help the plant if conditions are dry?

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Increased root production
§ allows the plant to explore more soil (volume and
depth) for extracting water
§ Reduction in leaf area
§ decreases the amount of solar radiation striking the
plant
§ decreases the amount of surface area that is losing
water through transpiration
§ Changes in leaf size and shape
§ leaves tend to be smaller and thicker

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ In combination, this increases the uptake of water
per unit leaf area, and reduces the total amount of
water lost by transpiration.

© 2016 Pearson Education, Ltd.

Figure 6.13
8

R(g):LA (cm2)
4

0
3 4 5 6 7
In water supply rate (ml/day)
(a)

0.7

% Allocation to leaves
0.6

0.5

0.4
3 4 5 6 7
In water supply rate (ml/day)
(b)

300

250
SLA (cm2/g)

200

150

100

50

0
3 4 5 6 7
In water supply rate (ml/day)
(c)
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Over long periods of time, modifications of
photosynthesis have evolved in some plants that
increase the efficiency of water use in warmer and
drier environments
§ Review – plants in areas without water limitation
generally use C3 photosynthesis
§ Mesophyll cells – location of light energy capture
and conversion, transformation of CO2 into sugars
§ Products of photosynthesis move into the vascular
bundles to be transported around the plant

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Plants that undergo C4 photosynthesis have a
different leaf anatomy
§ Two types of photosynthetic cells
§ mesophyll cells
§ bundle sheath cells – surround the bundles of
vascular tissue

© 2016 Pearson Education, Ltd.

Figure 6.14a

Mesophyll cell
Bundle sheath Photosynthetic
cell cells of C4
plant leaf

Vein
(vascular tissue)

Stoma
(a) C4 anatomy

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ In the mesophyll cells
§ CO2 reacts with PEP, a 3-carbon molecule
§ reaction catalyzed by PEP carboxylase
§ produces a 4-carbon molecule – OAA
§ OAA is rapidly transformed into a 4-carbon acid
(malic or aspartic), hence the name C4
§ Organic acids are transported to the bundle sheath
cells
§ Enzymes break them down to form CO2
§ This CO2 then enters the C3 pathway

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ What advantage does C4 photosynthesis provide?

© 2016 Pearson Education, Ltd.

Figure 6.14b

CO2
Mesophyll
cell
PEP
carboxylase Air space
Oxaloacetate (4 C) PEP (3 C)

Malate (4 C) ATP

Pyruvate (3 C)
CO2
Bundle
sheath
cell Calvin
cycle

Simple sugars

Vascular
tissue

(b) The C4 pathway

© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Advantages of C4 photosynthesis
§ PEP does not react with oxygen
§ photorespiration does not occur
§ conversion of organic acids into CO2 within the
bundle sheath cells concentrates CO2
§ reaches much higher concentrations than in C3
§ increases efficiency of CO2 and RuBP reaction
§ Spatially separates the steps in two different parts of
the leaf

© 2016 Pearson Education, Ltd.

Figure 6.16a

C4
CO2

Mesophyll
CO2 incorporated
cell Organic acid into four-carbon
organic acids
(carbon fixation)

Bundle
CO2
sheath
cell
Organic acids
Calvin release CO2 to
cycle Calvin cycle

Simple sugars

(a) Spatial separation of steps.

In C4 plants, carbon fixation and
the Calvin cycle occur in different
types of cells.
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ C4 plants generally have a higher maximum rate of
photosynthesis than C3 plants
§ Greater water-use efficiency is an advantage in hot,
dry climates
§ Increased energy to produce PEP and enzyme is
the cost
§ Most C4 plants are
§ grasses in tropical or subtropical environments
§ shrubs in arid or salty environments

© 2016 Pearson Education, Ltd.

Figure 6.15

20

40

60
80

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Hot deserts have more severe environments
§ high levels of solar radiation
§ eater availability is very low
§ Plants living in these conditions (mainly members of
the Cactaceae, Euphorbiaceae, and Crassulaceae
families) have a different type of photosynthesis
§ CAM pathway – crassulacean acid metabolism
§ Similar to the C4 pathway, but the steps are
separated temporally instead of spatially

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ CAM plants open stomata at night
§ take up CO2
§ convert to malic acid using PEP
§ large amounts of malic acid accumulate in mesophyll
cells
§ Close stomata during the day
§ malic acid converted to CO2
§ C3 pathway used to fix CO2 and produce sugars
§ How is this adaptive for plants living in a hot, dry
environment?

© 2016 Pearson Education, Ltd.

 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ CAM plants dramatically reduce water loss by
opening stomata only at night

© 2016 Pearson Education, Ltd.

Figure 6.16b

CAM CO
2

Night
CO2 incorporated
Organic acid
into four-carbon
organic acids
(carbon fixation)

Day CO2

Organic acids
Calvin release CO2 to
cycle Calvin cycle

Simple sugars

(b) Temporal separation of steps.

In CAM plants, carbon fixation
and the Calvin cycle occur in the
same cells at different times.
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Other adaptations seen in plants adapted to different
soil moisture environments
§ Plants in dry environments generally have lower
stomatal conductance
§ fewer and smaller stomata
§ increased water-use efficiency
§ decreased rate of photosynthesis
§ Greater allocation of carbon to produce roots
instead of shoots and leaves
§ increases area for extracting water (volume and
depth)
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ This is a trade-off because reduced leaf area
decreases the gain of carbon from photosynthesis

© 2016 Pearson Education, Ltd.

Figure 6.17
1.2
Forest and woodland

1.0

0.8

Root(g): shoot(g)
0.6

0.4

0.2

0.0
0 1000 2000 3000 4000
(a)
30
Shrubs and grassland

25

20
Root(g): shoot(g)

15

10

0
0 1000 2000 3000 4000
Precipitation (mm/yr)
(b)
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ Structural features can reduce the amount of energy
striking the leaf, increase loss of heat through
convection, reduce the loss of water through
transpiration
§ leaves of plants in dry environments are generally
smaller and thicker (lower SLA)
§ some have leaves covered with hairs that scatter
solar radiation
§ some have leaves coated with waxes and resins that
reflect light and reduce absorption

© 2016 Pearson Education, Ltd.

Figure 6.18

Leaf mass per unit leaf area (g/m2)

Eucalyptus miniata
300
Eucalyptus tetrodonta
Eucalyptus tectifica
Corymbia latifolia
250 Corymbia terminalis
Eucalyptus pruinosa
Eucalyptus coolabah
Corymbia aparrerinja
200

150

0 500 1000 1500 2000

Mean annual precipitation (mm)
(a)

120
Average leaf width (mm)

100

80

60
S. South America
40
N. South America
Australia
20
Central America
0
0 1000 2000 3000 4000
Mean annual precipitation (mm)
(b)
© 2016 Pearson Education, Ltd.
 Section 6.9 The Link between Water Demand
and Temperature Influences Plant Adaptations
§ In areas that have a wet and dry season some
plants drop their leaves when the dry season begins
§ drought deciduous
§ new leaves grow just before the rainy season begins

© 2016 Pearson Education, Ltd.

 Section 6.10 Plants Exhibit Both Acclimation and
Adaptation in Response to Variations in Environmental
Temperatures
§ Plants are sessile organisms
§ Exposed to wide variations in temperature on many
spatial scales and time scales
§ All plants experience daily changes in temperature
§ Where do plants experience seasonal changes in
temperature?

© 2016 Pearson Education, Ltd.

 Section 6.10 Plants Exhibit Both Acclimation and
Adaptation in Response to Variations in Environmental
Temperatures
§ Plants at higher latitudes and those living farther
from large bodies of water experience greater
seasonal changes in temperature
§ Tmin, Topt, and Tmax are usually
§ lower in species adapted to cooler environments
§ higher in species adapted to warmer environments

© 2016 Pearson Education, Ltd.

Figure 6.19

% of maximum photosynthesis

100

80

60

40 Neuropogon
Ambrosia
Tidestromia
20
Atriplex

0
-5 0 5 10 15 20 25 30 35 40
Leaf temperature (°C)

© 2016 Pearson Education, Ltd.

 Section 6.10 Plants Exhibit Both Acclimation and
Adaptation in Response to Variations in Environmental
Temperatures
§ C4 plants live in warmer, drier environments and
have higher optimal temperatures for photosynthesis
than C3
§ Topt is higher for PEP carboxylase than rubisco

© 2016 Pearson Education, Ltd.

Figure 6.20

1.6

(c)
Photosynthesis (mg CO2/m2/s)

1.2

0.8

0.4 (a)
(b)

0 10 20 30 40
Temperature (°C)

© 2016 Pearson Education, Ltd.

 Section 6.10 Plants Exhibit Both Acclimation and
Adaptation in Response to Variations in Environmental
Temperatures
§ Plants can acclimate to different thermal conditions
§ When individuals of the same species are grown at
different temperatures in the laboratory or
greenhouse, divergence in temperature response is
often observed:
§ Topt lower for plants grown under cooler conditions
§ Topt higher for plants grown under warmer conditions
§ Same shift in temperature response is observed in
response to seasonal temperature changes

© 2016 Pearson Education, Ltd.

Figure 6.21

40

23°C day/18°C night

CO2 uptake (µmol/m2/s)

30

20

43°C day/30°C night

10

0
10 20 30 40 50
Temperature (°C)
© 2016 Pearson Education, Ltd.
Figure 6.22

Net photosynthesis (µmol CO2/m2/s) 30

Spreading creosote bush
Topt Larrea divaricata

25 Topt
January

20
September

15
10 20 30 40
Temperature (°C)
© 2016 Pearson Education, Ltd.
 Section 6.10 Plants Exhibit Both Acclimation and
Adaptation in Response to Variations in Environmental
Temperatures
§ Periods of extreme heat or cold can damage plant
cells and tissues
§ Cold-tolerant plants show adaptations
§ frost hardening – the genetic ability to tolerate
extreme cold; variable within and among species
§ produce compounds that allow leaves to survive
freezing temperatures
§ needle leaf evergreens – pine and spruce
§ some species are winter deciduous – shed their
leaves before the beginning of the cold season

© 2016 Pearson Education, Ltd.

 Section 6.10 Plants Exhibit Both Acclimation and
Adaptation in Response to Variations in Environmental
Temperatures
§ Plants display many physiological and
morphological mechanisms that allow them to adjust
to prevailing environmental temperatures
§ There are trade-offs between performance at higher
temperatures versus lower temperatures
§ Shift in Tmin is usually associated with a shift in Tmax
§ shifts in enzymes and membranes (acclimation and
adaptation) to lower temperatures are usually
maladaptive at higher temperatures

© 2016 Pearson Education, Ltd.

 Section 6.11 Plants Exhibit Adaptations to
Variations in Nutrient Availability
§ Plants must have a variety of chemical elements for
metabolic processes and synthesis of new tissues
(listed in Table 6.1)
§ What is the difference between a macronutrient and
a micronutrient?
§ How do plants acquire these nutrients?

© 2016 Pearson Education, Ltd.

 Section 6.11 Plants Exhibit Adaptations to
Variations in Nutrient Availability
§ Macronutrients are nutrients that are needed in large
amounts
§ carbon, hydrogen, oxygen
§ derived from CO2 and H2O
§ nitrogen, phosphorus, potassium, calcium,
magnesium, sulfur
§ terrestrial plants acquire from the soil
§ aquatic autotrophs acquire from the substrate or water
§ Micronutrients (trace elements) are nutrients that are
needed in smaller amounts

© 2016 Pearson Education, Ltd.

 Section 6.11 Plants Exhibit Adaptations to
Variations in Nutrient Availability
§ Rate of nutrient absorption depends on nutrient
concentrations in the soil or water
§ When concentration at the root surface declines, the
rate of absorption declines
§ Leads to reduction in tissue nutrient concentration
§ a decrease in nitrogen concentrations in the leaf
reduces the maximum rate of photosynthesis
§ reduces production of rubisco and cholorphyll
§ How do plants respond to reduced nutrient
availability?

© 2016 Pearson Education, Ltd.

Figure 6.24

Root weight (g)/ Shoot (leaf + stem) weight (g)

1.0

Silybum marianum
0.8

0.6

0.4

0.2

0
1/64 1/16 1/4 1 4 16
Relative nutrient concentration
© 2016 Pearson Education, Ltd.
 Section 6.11 Plants Exhibit Adaptations to
Variations in Nutrient Availability
§ If nutrient availability is reduced, plants allocate
carbon to root growth instead of shoot growth
§ Phenotypic plasticity
§ compensates for decrease in nutrient absorption per
unit root by increasing the root area
§ increases the soil volume from which nutrients are
absorbed
§ reduces allocation to leaves, photosynthesis
decreases, growth rate is reduced
§ How do plants live in low-nutrient environments?

© 2016 Pearson Education, Ltd.

 Section 6.11 Plants Exhibit Adaptations to
Variations in Nutrient Availability
§ Plants in low-nutrient environments possess a
number of adaptations
§ a low absorption rate – absorb fewer nutrients in
fertile soils, but higher levels when nutrients are
limited
§ higher ratio of roots to shoots
§ lower growth rate
§ increased leaf longevity

© 2016 Pearson Education, Ltd.

Figure 6.26

100

Leaf nitrogen (mg/g)

10

1 10 100 1000
(a) Leaf life span (months)

1000
Net photosynthesis (nmol/g/s)

100

10

1 10 100 1000
(b) Leaf life span (months)
© 2016 Pearson Education, Ltd.
 Agrostis stolonifera and A. canina are both species of
bent grass. This graph shows the response of each to
an experiment in which the plants were grown at
different levels of nitrogen fertilizer (1 ppm to 243
ppm). Based on these results, which of the following is
true?

A. Both species normally grow in nutrient-rich soil.

B. A. stolonifera normally grows in more nutrient-poor
soil.
C. Both species normally grow in nutrient-poor soil.
D. A. canina normally grows in more nutrient-poor soil.

© 2016 Pearson Education, Ltd.

Figure 6.25

5.0 Agrostis stolonifera

4.0
3.0
2.0
Plant weight (g)

1.0

0.4
Agrostis canina

0.2

0
1 3 9 27 81 243
Nitrogen (ppm)

© 2016 Pearson Education, Ltd.

 Agrostis stolonifera and A. canina are both species of
bent grass. This graph shows the response of each to
an experiment in which the plants were grown at
different levels of nitrogen fertilizer (1 ppm to 243
ppm). Based on these results, which of the following is
true?

A. Both species normally grow in nutrient-rich soil.

B. A. stolonifera normally grows in more nutrient-poor
soil.
C. Both species normally grow in nutrient-poor soil.
D. A. canina normally grows in more nutrient-poor
soil.

© 2016 Pearson Education, Ltd.

 Section 6.12 Plant Adaptations to the Environment
Reflect a Trade-off between Growth Rate and Tolerance

§ Plant adaptations to abiotic environmental

conditions show a trade-off between characteristics
that allow high rates of photosynthesis and growth in
resource rich environments and the ability to tolerate
resource poor environments
§ What are some examples of these trade-offs?

© 2016 Pearson Education, Ltd.

 Section 6.12 Plant Adaptations to the Environment
Reflect a Trade-off between Growth Rate and Tolerance

§ Optimal photosynthesis requires:

§ warm temperatures
§ adequate supplies of light, water, mineral nutrients
§ What happens when these resources change?

© 2016 Pearson Education, Ltd.

 Section 6.12 Plant Adaptations to the Environment
Reflect a Trade-off between Growth Rate and Tolerance

§ Plants that are adapted to low-light conditions:

§ have lower rates of respiration
§ have lower light compensation point
§ maintain a positive rate of photosynthesis
§ under high-light levels, growth is limited
§ Plants that are adapted to dry conditions:
§ have an increased root/shoot ratio
§ have a reduced leaf size and area
§ under moist conditions, growth is limited

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants
Respond to Increasing Atmospheric CO2
§ What is the current atmospheric concentration of
CO2?
§ How can increasing concentrations of atmospheric
CO2 affect plants?

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ Review
§ What metabolic process in plants uses CO2?
§ How does CO2 enter plant leaves?

§ How would an increase in CO2 concentrations affect

this metabolic process in plants?

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ The CO2 fertilization effect
§ as the level of CO2 increases in the atmosphere, the
diffusion gradient between the air and the interior of
the leaf increases
§ more CO2 moves into the leaf
§ increased CO2 concentration in mesophyll cells with
increase the rate of photosynthesis
§ will also reduce photorespiration
§ overall, enhances the potential rates of new
photosynthesis

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ The same effect is not seen in C4 plants

© 2016 Pearson Education, Ltd.

Figure 6.28

C3

30
Photosynthesis (µmol/m2/s)

C4

20

10

Current 2´
CO2 levels CO2
0
0 200 400 600 800 1000
CO2 (ppm)
© 2016 Pearson Education, Ltd.
 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ Plants also show a reduction in stomatal
conductance when CO2 levels are elevated
§ conductance includes the number of stomata per area
and the size of the opening
§ Why does this happen?

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ Short term – the aperture size is reduced (the
stomata partially close)
§ Longer term – stomata density decreases
(developmental plasticity)
§ These changes
§ reduce the rate of transpiration
§ increase water use efficiency

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ Most experiments have been done in controlled
environmental conditions
§ How does increasing CO2 level affect plants in open
air conditions (field experiments)?
§ FACE – Free-air CO2 enrichment experiments
§ Different species grown under 567 ppm CO2
§ across all species, stomatal conductance was
reduced by 22 percent
§ greater decrease in grasses and herbaceous crops,
less decrease in trees, shrubs, and forbs

© 2016 Pearson Education, Ltd.

Figure 6.30a

C4 (25)

C3 (267)

C4 crop (11)

C4 grass (13)
C3 crop (42)

C3 grass (24)

Forb (16)
Shrub (56)

Tree (98)
-40 -30 -20 -10 0
% Change at elevated CO2
(a)
© 2016 Pearson Education, Ltd.
 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ What about the effect on photosynthetic rate?
§ Much more variable across plant type and
environment
§ Rate in C3 plants up an average of 31 percent
§ trees showed the largest response to elevated CO2
§ shrubs showed the smallest response
§ Rate also increased in C4 plants
§ indirect effect of reduced stomatal conductance so
transpiration was reduced and soil water status
improved

© 2016 Pearson Education, Ltd.

Figure 6.30b

C4 crop (18)

C4 grass (11)

C3 crop (51)

C3 grass (62)

Shrub (19)

Tree (191)

-10 0 10 20 30 40 50 60
% Change at elevated CO2
(b)

© 2016 Pearson Education, Ltd.

 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ What about the effect of long-term exposure to
elevated CO2 levels?
§ Poorter and Pérez-Soba reviewed results from more
than 600 experiments looking at plant growth under
elevated CO2 conditions
§ All three photosynthetic pathways represented
§ Average increase in biomass was reported:
§ C3 plants – strongest response – 47 percent
§ CAM plants – based on six species – 21 percent
§ C4 plants – 11 percent

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Figure 6.32

2.5

49%
41%
59%
2.0
BER

1.5 11%
21%

1.0

0.5
Crop Wild Woody C4 CAM
C3 species
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 Ecological Issues & Applications: Plants Respond
to Increasing Atmospheric CO2
§ Do these enhanced effects on plant growth persist?
§ Longer-term studies suggest that the effects are
short-lived
§ some plants downregulate – produce less rubisco at
elevated CO2 levels
§ photosynthesis rate declines to previous level
§ some studies show plants reallocate carbon, produce
fewer leave and more roots
§ Nitrogen availability has been shown to influence the
response of plants to elevated CO2

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