Contents

Chapter 29. Resource Acquisition, Nutrition, and Transport in Vascular Plants

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Key Steps in the Evolution of Vascular Plants

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Shoot Architecture and Light Capture
• Plant success is generally related to photosynthesis
• Natural selection has resulted in many shoot adaptations to acquire light
and CO2 more efficiently

• Stems serve as conduits for water and nutrients and as supporting

structures for leaves
• Stem lengths, widths, and branching patterns affect light capture
• There is an energy trade-off between growing tall to avoid shading and
branching to harvest light

• The arrangement of leaves on a stem is important to light capture

• The leaves of many plants are arranged in an ascending spiral with each
successive leaf emerging 137.5° from the site of the previous one
• This pattern may reduce shading of lower leaves
• Shaded, nonproductive leaves are shed in a process called self-pruning

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Shoot Architecture and Light Capture

• Leaf orientation affects light absorption

• In low-light conditions, horizontal leaves capture more sunlight
• In sunny conditions, vertical leaves are less damaged by sun and allow
light to reach lower leaves 4
The Apoplast and Symplast: Transport Continuums
• The apoplast consists of everything external to the plasma membrane of
living cells
• The symplast consists of the cytosol of all the living cells in a plant, as
well as the plasmodesmata

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Short-Distance Transport of Solutes Across Plasma Membranes

• Selective permeability of the plasma membrane controls the movement

of substances into and out of cells
• Both active and passive transport occur in plants

• In plants, membrane potential is established through pumping H+ by

proton pumps
• Plant cells use the energy of H+ gradients to cotransport other solutes by
active transport; Na+ is typically cotransported in animals

• Osmosis, the diffusion of free water across a membrane, determines

the net uptake or water loss by a cell
• The direction of water flow is determined by water potential, a quantity
that includes the effects of solute concentration and pressure
• Solutions with higher solute concentrations have relatively lower water
potential
Ψ = ΨS + ΨP
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Water Movement Across Plant Cell Membranes
• Because they contain solutes, cells have lower water potential than pure
water
• If a flaccid cell is placed in pure water, water will move into the cell and
the cell will become turgid (firm)

• When turgid cells push against each other, the tissue stiffens
• Turgor loss in plants causes wilting, which can be reversed when the
plant is watered

• Water molecules move across plasma membranes through transport

proteins called aquaporins
• Aquaporin channels can open or close to change the rate of water
movement across the membrane 7
Water Movement Across Plant Cell Membranes

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Macronutrients and Micronutrients
• Nine of the essential elements are called macronutrients because
plants require them in relatively large amounts
• The remaining essential elements, called micronutrients, are only
required in tiny quantities
• Micronutrients include chlorine, iron, manganese, boron, zinc, copper,
nickel, and molybdenum

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Relationships with Other Organisms

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Bacteria in the Nitrogen Cycle
• Globally, nitrogen is the nutrient most limiting to plant growth
• Plants can only absorb nitrogen as nitrate (NO3-) or ammonium (NH4+)
• The nitrogen cycle describes transformations of nitrogen and nitrogenous
compounds in nature
• Most usable soil nitrogen comes from actions of soil bacteria

• Conversion to NH4+
• Ammonifying bacteria break down organic compounds and release ammonium
(NH4+)
– Nitrogen-fixing bacteria convert N2 gas into NH3
– NH3 combines with H+ from soil to form NH4+
• Conversion to NO3-
• Nitrifying bacteria oxidize NH4+ to nitrate (NO2-) and then oxidize nitrite to nitrate
(NO3-)
• Different nitrifying bacteria mediate each step

• Nitrogen is lost to the atmosphere when denitrifying bacteria convert NO3- to N

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Bacteria in the Nitrogen Cycle

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Nitrogen-Fixing Bacteria: A Closer Look
• Nitrogen (N2) is abundant in the atmosphere, but the triple bond between
the nitrogen atoms makes it unusable to plants
• Nitrogen fixation is the conversion of nitrogen from N2 to NH3 by
bacteria:

• Some nitrogen-fixing bacteria are free-living; others form intimate

associations with plant roots

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Epiphytes, Parasitic Plants, and Carnivorous Plants
• Some plants have nutritional adaptations that use other organisms in
nonmutualistic ways
• Three unusual adaptations are
– Epiphytes
– Parasitic plants
– Carnivorous plants

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Epiphytes, Parasitic Plants, and Carnivorous Plants
• Epiphytes grow on other plants and obtain water and minerals from rain,
rather than tapping their hosts for sustenance

• Parasitic plants absorb water, sugars, and minerals directly from their
living host plant
• Some species also photosynthesize, but others rely entirely on the host
plant for sustenance

• Carnivorous plants have adaptations for trapping insects and other

small animals
• They are photosynthetic but obtain nitrogen by killing and digesting prey

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Carnivorous Plants

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Transport of Water and Minerals into the Xylem

• Water and minerals can travel to the vascular cylinder through the cortex
via
1. The apoplastic route, along cell walls and extracellular spaces
2. The symplastic route, in the cytoplasm, moving between cells
through plasmodesmata
3. The transmembrane route, moving from cell to cell by crossing cell
membranes and cell walls

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Transport of Water and Minerals into the Xylem

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Transport of Water and Minerals into the Xylem
• The waxy Casparian strip of the endodermal wall blocks apoplastic
transfer of minerals and water from the cortex to the vascular cylinder
• Water and minerals must cross a selectively permeable plasma
membrane before entering the vascular cylinder

2020. PNAS
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Bulk Flow Transport via the Xylem
• The transport of xylem sap involves transpiration, the loss of water
vapor from leaves and other parts of the shoot
• Unless transpired water is replaced by water traveling up from the roots,
the leaves will wilt

• According to the cohesion-tension hypothesis, transpiration provides

the pull for the ascent of xylem sap
• Cohesion between water molecules transmits the pull along the xylem
from shoots to roots
• Transpirational pull is generated when water vapor in the air spaces of
a leaf diffuses down its water potential gradient and exits via stomata
• This negative pressure is transferred to the xylem and water is pulled
from the xylem into the leaf
• The transpirational pull on xylem sap is transmitted from leaves to root
tips and into the soil solution
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Pulling Xylem Sap: The Cohesion-Tension Hypothesis

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Pulling Xylem Sap: The Cohesion-Tension Hypothesis
Cohesion and adhesion
in the ascent of xylem
sap
• Water molecules are
attracted to each other
through cohesion and to
the walls of the xylem
cells through adhesion
• Cohesion transmits the
pull down a column of
xylem sap; adhesion
helps to offset the force
of gravity

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Mechanisms of Stomatal Opening and Closing
• Guard cells open and close the stomata to help balance water
conservation with gas exchange
• Changes in turgor pressure open and close stomata
– When turgid, guard cells bow outward and the pore between them
opens
– When flaccid, guard cells become less bowed and the pore closes

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Stimuli for Stomatal Opening and Closing
• All eukaryotic organisms have internal clocks; circadian rhythms are 24-
hour cycles
• The internal clock in guard cells ensure that stomata continue their daily
rhythm of opening and closing even in a dark location

• Drought stress can cause stomata to close during the daytime

• The hormone abscisic acid (ABA) is produced in response to water
deficiency and causes the closure of stomata
• CO2 absorption is restricted when stomata are closed, causing
photosynthesis to slow

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Movement from Sugar Sources to Sugar Sinks
• The products of
photosynthesis are
transported through phloem
by the process of
translocation
• Phloem sap is an aqueous
solution, high in sucrose, that
travels from sugar sources to
sinks
• A sugar source is an organ
that is a net producer of
sugar, such as mature leaves
• A sugar sink is an organ that
is a net consumer or storer of
sugar, such as growing roots,
fruits, or leaves

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Movement from Sugar Sources to Sugar Sinks
• Sugar must be loaded into sieve-tube elements before being exported to sinks
• Sugar may move by symplastic or both symplastic and apoplastic pathways
• Companion cells can enhance solute movement between the apoplast and
symplast

• In most plants, active transport is required to move sucrose against its

concentration gradient from mesophyll cells to phloem cells
• Proton pumping and cotransport of sucrose and H+ enable phloem loading

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