Concept 37.

2: Plants require essential elements to

complete their life cycle

a) Soil, water, and air all contribute to plant growth

a)80–90% of a plant’s fresh mass is water
b)96% of a plant’s dry mass is from CO2 assimilated into
carbohydrates during photosynthesis
c)4% of a plant’s dry mass is inorganic substances from
soil

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Essential Elements

a)More than 50 chemical elements have been

identified among the inorganic substances in
plants, but not all of these are essential to plants
b)There are 17 essential elements, chemical
elements required for a plant to complete its life
cycle and reproduce
c) Researchers use hydroponic culture to
determine which chemical elements are essential

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Figure 37.7

Control: Solution Experimental: Solution

containing all minerals without potassium

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Table 37.1

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 a)Nine of the essential elements are called
macronutrients because plants require them in
relatively large amounts
b)The macronutrients are carbon, oxygen,
hydrogen, nitrogen, phosphorus, sulfur,
potassium, calcium, and magnesium

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 a)The remaining eight are called micronutrients
because plants need them in very small amounts
b)The micronutrients are chlorine, iron, manganese,
boron, zinc, copper, nickel, and molybdenum
c) Plants with C4 and CAM photosynthetic pathways
also need sodium
d)Micronutrients function as cofactors, nonprotein
helpers in enzymatic reactions

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Symptoms of Mineral Deficiency

a) Symptoms of mineral deficiency depend on the

nutrient’s function and mobility within the plant
b) Deficiency of a mobile nutrient usually affects older
organs more than young ones
c) Deficiency of a less mobile nutrient usually affects
younger organs more than older ones
d) The most common deficiencies are those of nitrogen,
potassium, and phosphorus

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Improving Plant Nutrition by Genetic Modification

a) Plants can be genetically engineered to better fit the

soil

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Resistance to Aluminum Toxicity

a)Aluminum in acidic soils damages roots and

greatly reduces crop yields
b)The introduction of bacterial genes into plant
genomes can cause plants to secrete acids that
bind to and tie up aluminum

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Smart Plants

a)“Smart” plants inform the grower of a nutrient

deficiency before damage has occurred
b)A blue tinge indicates when these plants need
phosphate-containing fertilizer

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Figure 37.8

No phosphorus Beginning Well-developed

deficiency phosphorus phosphorus
deficiency deficiency

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Concept 37.3: Plant nutrition often involves
relationships with other organisms

a) Plants and soil microbes have a mutualistic

relationship
a)Dead plants provide energy needed by soil-dwelling
microorganisms
b)Secretions from living roots support a wide variety of
microbes in the near-root environment

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Bacteria and Plant Nutrition

a)The layer of soil closely surrounding the plant’s

roots is the rhizosphere
b)Rhizobacteria are free-living bacteria that occupy
the rhizosphere
c)Endophytes are nonpathogenic bacteria that live
between the cells of host plant tissues

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 a) Endophytes and rhizobacteria depend on nutrients
secreted by plant cells and, in return, help to enhance
plant growth by
a)Producing chemicals that stimulate plant growth
b)Producing antibiotics that protect roots from disease
c)Absorbing toxic metals or increasing nutrient availability

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 a) The species composition of bacterial communities
living endophytically and in the rhizosphere vary
markedly

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Figure 37.9

Endophytes
Clayey soil
34% similar
Endophytes
Porous soil

Outside rhizosphere
Clayey soil

Inside rhizosphere
Clayey soil

Inside rhizosphere
Porous soil

80% similar Outside rhizosphere

Porous soil

0 20 40 60 80
Percent similarity of the Younger roots
Bacteria (green) on surface of “species” composition Older roots
root (fluorescent LM) of communities

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 Bacteria in the Nitrogen Cycle

a)Nitrogen can be an important limiting nutrient for

plant growth
b)Plants can absorb nitrogen as either NO3– or NH4
c) Most soil nitrogen comes from actions of soil
bacteria
d)The nitrogen cycle transforms nitrogen and
nitrogen-containing compounds

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Figure 37.10

ATMOSPHERE N2 ATMOSPHERE N2

Proteins from humus Nitrate and

SOIL
(dead organic material) nitrogenous
organic
Nitrogen-fixing compounds
Microbial
bacteria exported in
decomposition
N2 Amino xylem to
NH3 acids shoot system
Denitrifying
(ammonia) Ammonifying bacteria
bacteria NH4
H
SOIL (from soil) NH 
4 NO −
2 NO 3
−

(ammonium) Nitrifying (nitrate) Nitrifying (nitrate)

bacteria bacteria

Root

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 a) Conversion to NH4
a)Ammonifying bacteria break down organic compounds
and release ammonia (NH3)
b)Nitrogen-fixing bacteria convert N2 into NH3
c)NH3 is converted to NH4

b) Conversion to NO3
a)Nitrifying bacteria oxidize NH3 to nitrite (NO2) then
nitrite to nitrate (NO3)

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 a)Nitrogen is lost to the atmosphere when
denitrifying bacteria convert NO3– to N2

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Nitrogen-Fixing Bacteria: A Closer Look

a)Nitrogen is abundant in the atmosphere, but

unavailable to plants because of the triple bond
between atoms in N2
b)Nitrogen fixation is the conversion of nitrogen
from N2 to NH3

c) Symbiotic relationships with nitrogen-fixing

Rhizobium bacteria provide some plant species
(e.g., legumes) with a source of fixed nitrogen
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 a)Along a legume’s roots are swellings called
nodules, composed of plant cells “infected” by
nitrogen-fixing Rhizobium bacteria

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Figure 37.11

Nodules

Roots

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 a)Inside the root nodule, Rhizobium bacteria
assume a form called bacteroids, which are
contained within vesicles formed by the root cell

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 a)The plant obtains fixed nitrogen from Rhizobium,
and Rhizobium obtains sugar and an anaerobic
environment
b)Each legume species is associated with a
particular strain of Rhizobium
c) The development of a nitrogen-fixing root nodule
depends on chemical dialogue between
Rhizobium bacteria and root cells of their specific
plant hosts
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Figure 37.12
Rhizobium
bacteria Dividing cells
Infection
1 Chemical signals in root cortex
thread
attract bacteria
and an infection 2 Bacteroids form.
thread forms. Bacteroid
Nodule Infected
vascular root hair Dividing cells
tissue in pericycle

Bacteroid Root hair

Bacteroids sloughed
off

Developing
root nodule

3 Growth
Sclerenchyma continues and
cells a root nodule
forms.
5 The mature nodule
grows to be many Nodule 4 The nodule develops
times the diameter vascular vascular tissue.
Bacteroid tissue
of the root.
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Nitrogen Fixation and Agriculture

a) Crop rotation takes advantage of the agricultural

benefits of symbiotic nitrogen fixation
b) A nonlegume such as maize is planted one year, and
the next year a legume is planted to restore the
concentration of fixed nitrogen in the soil

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 a) Instead of being harvested, the legume crop is often
plowed under to decompose as “green manure”
b) Nonlegumes such as alder trees and certain tropical
grasses benefit from nitrogen-fixing bacteria
c) Rice paddies often contain an aquatic fern that has
mutualistic cyanobacteria that fix nitrogen

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Fungi and Plant Nutrition

a) Mycorrhizae are mutualistic associations of fungi and

roots
b) The fungus benefits from a steady supply of sugar
from the host plant
c) The host plant benefits because the fungus increases
the surface area for water uptake and mineral
absorption
d) Mycorrhizal fungi also secrete growth factors that
stimulate root growth and branching

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Mycorrhizae and Plant Evolution

a)Early land plants would have encountered harsh

conditions 400 to 500 million years ago
b)These plants lacked the ability to extract essential
nutrients from the soil, while fungi were unable to
produce carbohydrates
c) Mycorrhizal fungi may have helped plants
overcome the challenges of life on land

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The Two Main Types of Mycorrhizae

a) Mycorrhizal associations consist of two major types

a)Ectomycorrhizae
b)Arbuscular mycorrhizae

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Figure 37.13
Epidermis Cortex Mantle (fungal sheath)

Epidermal

(Colorized SEM)
cell

Endodermis

Fungal
hyphae
between
1.5 mm cortical
Mantle
cells (LM)
(fungal sheath) 50 μm
(a) Ectomycorrhizae

Epidermis Cortex
Cortical cell

Endodermis
Fungal Fungal
hyphae vesicle
Casparian

10 μm
strip
Root Arbuscules
hair
Plasma
membrane (LM)
(b) Arbuscular mycorrhizae
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 a)In ectomycorrhizae, the mycelium of the fungus
forms a dense sheath over the surface of the root
b)These hyphae form a network in the apoplast, but
do not penetrate the root cells
c) Ectomycorrhizae occur in about 10% of plant
families including pine, oak, birch, and eucalyptus

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 a)In arbuscular mycorrhizae, microscopic fungal
hyphae extend into the root
b)These mycorrhizae penetrate the cell wall but not
the plasma membrane
c) Hyphae form branched arbuscules within cells;
these are important sites of nutrient transfer
d)Arbuscular mycorrhizae occur in about 85% of
plant species, including grains and legumes

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Agricultural and Ecological Importance of Mycorrhizae

a) Farmers and foresters often inoculate seeds with

fungal spores to promote formation of mycorrhizae
b) Some invasive exotic plants disrupt interactions
between native plants and their mycorrhizal fungi
a)For example, garlic mustard slows growth of other
plants by preventing the growth of mycorrhizal fungi

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Epiphytes, Parasitic Plants, and Carnivorous Plants

a) Some plants have nutritional adaptations that use

other organisms in nonmutualistic ways
b) Three unusual adaptations are
a)Epiphytes
b)Parasitic plants
c)Carnivorous plants

c) An epiphyte grows on another plant and obtains

water and minerals from rain

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Figure 37.14a

Staghorn fern, an epiphyte

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Figure 37.14b

Parasitic plants

Rafflesia, a non- Indian pipe, a non-

photosynthetic photosynthetic
parasite parasite of
mycorrhizae
Mistletoe, a photosynthetic parasite

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Figure 37.14c

Carnivorous plants

Sundew

Pitcher plants Venus flytraps

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 a) Parasitic plants absorb sugars and minerals from
their living host plant
b) Carnivorous plants are photosynthetic but obtain
nitrogen by killing and digesting mostly insects

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