1. Soil and Plant Nutrition
Chapter 37
Soil and Plant Nutrition

2. Overview: A Horrifying Discovery
Carnivory by pitcher plants is well-documented
An extreme example is Nepenthes rajah, a pitcher plant large enough to catch a rat
N. Rajah lives in very unproductive soil and uses carnivory to obtain nutrients such as calcium, potassium, and phosphorus
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3. Figure 37.1
Figure 37.1 A rat trap?

4. Concept 37.1: Soil contains a living, complex ecosystem
Plants obtain most of their water and minerals from the upper layers of soil
Living organisms play an important role in these soil layers
This complex ecosystem is fragile
The basic physical properties of soil are
Texture
Composition
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5. Soil Texture
Soil particles are classified by size; from largest to smallest they are called sand, silt, and clay
Soil is stratified into layers called soil horizons
Topsoil consists of mineral particles, living organisms, and humus, the decaying organic material
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6. Figure 37.2
A horizon
B horizon
Figure 37.2 Soil horizons.
C horizon

7. The film of loosely bound water is usually available to plants
Soil solution consists of water and dissolved minerals in the pores between soil particles
After a heavy rainfall, water drains from the larger spaces in the soil, but smaller spaces retain water because of its attraction to clay and other particles
The film of loosely bound water is usually available to plants
Loams are the most fertile topsoils and contain equal amounts of sand, silt, and clay
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8. Topsoil Composition
A soil’s composition refers to its inorganic (mineral) and organic chemical components
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9. Inorganic Components
Cations (for example K+, Ca2+, Mg2+) adhere to negatively charged soil particles; this prevents them from leaching out of the soil through percolating groundwater
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10. During cation exchange, cations are displaced from soil particles by other cations
Displaced cations enter the soil solution and can be taken up by plant roots
Negatively charged ions do not bind with soil particles and can be lost from the soil by leaching
Animation: How Plants Obtain Minerals from Soil
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11. Soil particle   K K        Ca2 Ca2 Mg2 K H H2O  CO2
Figure 37.3
Soil particle   K K       
Ca2
Ca2
Mg2 K H
H2O  CO2
H2CO3
HCO3  H
Figure 37.3 Cation exchange in soil.
Root hair
Cell wall

12. Organic Components
Humus builds a crumbly soil that retains water but is still porous
It also increases the soil’s capacity to exchange cations and serves as a reservoir of mineral nutrients
Topsoil contains bacteria, fungi, algae, other protists, insects, earthworms, nematodes, and plant roots
These organisms help to decompose organic material and mix the soil
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13. Soil Conservation and Sustainable Agriculture
Soil management, by fertilization and other practices, allowed for agriculture and cities
In contrast with natural ecosystems, agriculture depletes the mineral content of soil, taxes water reserves, and encourages erosion
The American Dust Bowl of the 1930s resulted from soil mismanagement
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14. Figure 37.4
Figure 37.4 A massive dust storm in the American Dust Bowl during the 1930s.

15. At present, 30% of the world’s farmland has reduced productivity because of soil mismanagement
The goal of sustainable agriculture is to use farming methods that are conservation-minded, environmentally safe, and profitable
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16. Irrigation
Irrigation is a huge drain on water resources when used for farming in arid regions
For example, 75% of global freshwater use is devoted to agriculture
The primary source of irrigation water is underground water reserves called aquifers
The depleting of aquifers can result in land subsidence, the settling or sinking of land
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17. Figure 37.5
Figure 37.5 Sudden land subsidence.

18. Drip irrigation requires less water and reduces salinization
Irrigation can lead to salinization, the concentration of salts in soil as water evaporates
Drip irrigation requires less water and reduces salinization
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19. Fertilization
Soils can become depleted of nutrients as plants and the nutrients they contain are harvested
Fertilization replaces mineral nutrients that have been lost from the soil
Commercial fertilizers are enriched in nitrogen (N), phosphorus (P), and potassium (K)
Excess minerals are often leached from the soil and can cause algal blooms in lakes
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20. Organic fertilizers are composed of manure, fishmeal, or compost
They release N, P, and K as they decompose
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21. Adjusting Soil pH
Soil pH affects cation exchange and the chemical form of minerals
Cations are more available in slightly acidic soil, as H+ ions displace mineral cations from clay particles
The availability of different minerals varies with pH
For example, at pH 8 plants can absorb calcium but not iron
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22. Controlling Erosion
Topsoil from thousands of acres of farmland is lost to water and wind erosion each year in the United States
Erosion of soil causes loss of nutrients
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23. Erosion can be reduced by
Planting trees as windbreaks
Terracing hillside crops
Cultivating in a contour pattern
Practicing no-till agriculture
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24. Figure 37.6
Figure 37.6 Contour tillage.

25. Phytoremediation
Some areas are unfit for agriculture because of contamination of soil or groundwater with toxic pollutants
Phytoremediation is a biological, nondestructive technology that reclaims contaminated areas
Plants capable of extracting soil pollutants are grown and are then disposed of safely
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26. Concept 37.2: Plants require essential elements to complete their life cycle
Soil, water, and air all contribute to plant growth
80–90% of a plant’s fresh mass is water
4% of a plant’s dry mass is inorganic substances from soil
96% of plant’s dry mass is from CO2 assimilated during photosynthesis
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27. Macronutrients and Micronutrients
More than 50 chemical elements have been identified among the inorganic substances in plants, but not all of these are essential to plants
There are 17 essential elements, chemical elements required for a plant to complete its life cycle
Researchers use hydroponic culture to determine which chemical elements are essential
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28. TECHNIQUE Control: Solution containing all minerals
Figure 37.7
TECHNIQUE
Figure 37.7 Research Method: Hydroponic Culture
Control: Solution containing all minerals
Experimental: Solution without potassium

29. Table 37.1
Table 37.1 Essential Elements in Plants

30. Nine of the essential elements are called macronutrients because plants require them in relatively large amounts
The macronutrients are carbon, oxygen, hydrogen, nitrogen, phosphorus, sulfur, potassium, calcium, and magnesium
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31. Plants with C4 and CAM photosynthetic pathways also need sodium
The remaining eight are called micronutrients because plants need them in very small amounts
The micronutrients are chlorine, iron, manganese, boron, zinc, copper, nickel, and molybdenum
Plants with C4 and CAM photosynthetic pathways also need sodium
Micronutrients function as cofactors, nonprotein helpers in enzymatic reactions
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32. Symptoms of Mineral Deficiency
Symptoms of mineral deficiency depend on the nutrient’s function and mobility within the plant
Deficiency of a mobile nutrient usually affects older organs more than young ones
Deficiency of a less mobile nutrient usually affects younger organs more than older ones
The most common deficiencies are those of nitrogen, potassium, and phosphorus
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33. Healthy Phosphate-deficient Potassium-deficient Nitrogen-deficient
Figure 37.8
Healthy
Phosphate-deficient
Potassium-deficient
Figure 37.8 The most common mineral deficiencies, as seen in maize leaves.
Nitrogen-deficient

34. Improving Plant Nutrition by Genetic Modification: Some Examples
Plants can be genetically engineered to better fit the soil
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35. Resistance to Aluminum Toxicity
Aluminum in acidic soils damages roots and greatly reduces crop yields
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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36. Flood Tolerance
Waterlogged soils deprive roots of oxygen and cause buildup of ethanol and toxins
The gene Submergence 1A-1 is responsible for submergence tolerance in flood-resistant rice
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37. Smart Plants
“Smart” plants inform the grower of a nutrient deficiency before damage has occurred
A blue tinge indicates when these plants need phosphate-containing fertilizer
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38. No phosphorus deficiency Beginning phosphorus deficiency
Figure 37.9
No phosphorus deficiency
Beginning phosphorus deficiency
Well-developed phosphorus deficiency
Figure 37.9 Deficiency warnings from “smart” plants.

39. No phosphorus deficiency
Figure 37.9a
Figure 37.9 Deficiency warnings from “smart” plants.
No phosphorus deficiency

40. Beginning phosphorus deficiency
Figure 37.9b
Beginning phosphorus deficiency
Figure 37.9 Deficiency warnings from “smart” plants.

41. Well-developed phosphorus deficiency
Figure 37.9c
Well-developed phosphorus deficiency
Figure 37.9 Deficiency warnings from “smart” plants.

42. Concept 37.3: Plant nutrition often involves relationships with other organisms
Plants and soil microbes have a mutualistic relationship
Dead plants provide energy needed by soil-dwelling microorganisms
Secretions from living roots support a wide variety of microbes in the near-root environment
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43. Soil Bacteria and Plant Nutrition
The layer of soil bound to the plant’s roots is the rhizosphere
The rhizosphere contains bacteria that act as decomposers and nitrogen-fixers
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44. Rhizobacteria
Free-living rhizobacteria thrive in the rhizosphere, and some can enter roots
The rhizosphere has high microbial activity because of sugars, amino acids, and organic acids secreted by roots
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45. Rhizobacteria can play several roles
Produce hormones that stimulate plant growth
Produce antibiotics that protect roots from disease
Absorb toxic metals or make nutrients more available to roots
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46. Bacteria in the Nitrogen Cycle
Nitrogen can be an important limiting nutrient for plant growth
The nitrogen cycle transforms nitrogen and nitrogen-containing compounds
Plants can absorb nitrogen as either NO3– or NH4
Most soil nitrogen comes from actions of soil bacteria
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47. Nitrogen-fixing bacteria Denitrifying bacteria
Figure 37.10
ATMOSPHERE N2 N2
ATMOSPHERE
SOIL
Nitrate and nitrogenous organic compounds exported in xylem to shoot system
Nitrogen-fixing bacteria N2
Denitrifying bacteria
H (from soil)
NH4
SOIL
NH3 (ammonia)
NH4 (ammonium)
NO3 (nitrate)
Ammonifying bacteria
Nitrifying bacteria
Figure The roles of soil bacteria in the nitrogen nutrition of plants.
Organic material (humus)
Root

48. Nitrogen-fixing bacteria
Figure 37.10a-1 N2
Nitrogen-fixing bacteria
Figure The roles of soil bacteria in the nitrogen nutrition of plants.
NH3 (ammonia)
Ammonifying bacteria
Organic material (humus)

49. Nitrogen-fixing bacteria Denitrifying bacteria
Figure 37.10a-2 N2 N2
ATMOSPHERE
SOIL
Nitrate and nitrogenous organic compounds exported in xylem to shoot system
Nitrogen-fixing bacteria
Denitrifying bacteria
H (from soil)
NH4
NH3 (ammonia)
NH4 (ammonium)
NO3 (nitrate)
Figure The roles of soil bacteria in the nitrogen nutrition of plants.
Ammonifying bacteria
Nitrifying bacteria
Organic material (humus)
Root

50. Conversion to NH4 Conversion to NO3–
Ammonifying bacteria break down organic compounds and release ammonia (NH3)
Nitrogen-fixing bacteria convert N2 into NH3
NH3 is converted to NH4
Conversion to NO3–
Nitrifying bacteria oxidize NH3 to nitrite (NO2–) then nitrite to nitrate (NO3–)
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51. Nitrogen is lost to the atmosphere when denitrifying bacteria convert NO3– to N2
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52. Nitrogen-Fixing Bacteria: A Closer Look
Nitrogen is abundant in the atmosphere, but unavailable to plants because of the triple bond between atoms in N2
Nitrogen fixation is the conversion of nitrogen from N2 to NH3
N2  8e  8 H  16 ATP  2 NH3  H2  16 ADP  16 Pi
Symbiotic relationships with nitrogen-fixing Rhizobium bacteria provide some plant species (e.g., legumes) with a source of fixed nitrogen
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53. Along a legume’s roots are swellings called nodules, composed of plant cells “infected” by nitrogen-fixing Rhizobium bacteria
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54. Bacteroids within vesicle Nodules
Figure 37.11
Bacteroids within vesicle
Nodules
Roots
5 m
Figure Root nodules on a legume.
(a) Soybean root
(b)
Bacteroids in a soybean root nodule

55. Nodules Roots (a) Soybean root Figure 37.11a
Figure Root nodules on a legume.
Roots
(a) Soybean root

56. 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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57. Bacteroids within vesicle
Figure 37.11b
Bacteroids within vesicle
Figure Root nodules on a legume.
5 m
(b)
Bacteroids in a soybean root nodule

58. The plant obtains fixed nitrogen from Rhizobium, and Rhizobium obtains sugar and an anaerobic environment
Each legume species is associated with a particular strain of Rhizobium
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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59. Dividing cells in root cortex 1
Figure 37.12
Rhizobium bacteria
Infection thread
Dividing cells in root cortex 1
Chemical signals attract bacteria and an infection thread forms. 2
Bacteroids form.
Infected root hair
Nodule vascular tissue
Bacteroid
Dividing cells in pericycle
Bacteroid
Bacteroids
Root hair sloughed off
Developing root nodule
Figure Development of a soybean root nodule. 3
Growth continues and a root nodule
forms.
Sclerenchyma cells 5
The mature nodule grows to be many times the diameter of the root. 4
Nodule vascular tissue
The nodule develops vascular tissue.
Bacteroid

60. Nitrogen Fixation and Agriculture
Crop rotation takes advantage of the agricultural benefits of symbiotic nitrogen fixation
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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61. Instead of being harvested, the legume crop is often plowed under to decompose as “green manure”
Nonlegumes such as alder trees and certain tropical grasses benefit from nitrogen-fixing bacteria
Rice paddies often contain an aquatic fern that has mutualistic cyanobacteria that fix nitrogen
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62. Fungi and Plant Nutrition
Mycorrhizae are mutualistic associations of fungi and roots
The fungus benefits from a steady supply of sugar from the host plant
The host plant benefits because the fungus increases the surface area for water uptake and mineral absorption
Mycorrhizal fungi also secrete growth factors that stimulate root growth and branching
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63. Mycorrhizae and Plant Evolution
Mycorrhizal fungi date to 460 million years ago and might have helped plants colonize land
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64. The Two Main Types of Mycorrhizae
Mycorrhizal associations consist of two major types
Ectomycorrhizae
Arbuscular mycorrhizae
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65. Mantle (fungal sheath)
Figure 37.13
Epidermis
Cortex
Mantle (fungal sheath)
Epidermal cell
(Colorized SEM)
Endodermis
Fungal hyphae between cortical cells
1.5 mm
Mantle (fungal sheath)
(LM)
50 m
(a) Ectomycorrhizae
Cortical cell
Epidermis
Cortex
Endodermis
Figure Mycorrhizae.
Fungal vesicle
Fungal hyphae
Casparian strip
Root hair
10 m
Arbuscules
Plasma membrane
(LM)
(b)
Arbuscular mycorrhizae (endomycorrhizae)

66. In ectomycorrhizae, the mycelium of the fungus forms a dense sheath over the surface of the root
These hyphae form a network in the apoplast, but do not penetrate the root cells
Ectomycorrhizae occur in about 10% of plant families including pine, spruce, oak, walnut, birch, willow, and eucalyptus
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67. Mantle (fungal sheath)
Figure 37.13aa
Epidermis
Cortex
Mantle (fungal sheath)
Epidermal cell
(Colorized SEM)
Endodermis
Fungal hyphae between cortical cells
1.5 mm
Mantle (fungal sheath)
Figure Mycorrhizae.
(LM)
50 m
(a) Ectomycorrhizae

68. Mantle (fungal sheath)
Figure 37.13ab
(Colorized SEM)
Figure Mycorrhizae.
1.5 mm
Mantle (fungal sheath)

69. Fungal hyphae between cortical cells
Figure 37.13ac
Epidermal cell
Fungal hyphae between cortical cells
Figure Mycorrhizae.
(LM)
50 m

70. In arbuscular mycorrhizae, microscopic fungal hyphae extend into the root
These mycorrhizae penetrate the cell wall but not the plasma membrane to form branched arbuscules within root cells
Hyphae can form arbuscules within cells; these are important sites of nutrient transfer
Arbuscular mycorrhizae occur in about 85% of plant species, including grains and legumes
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71. Arbuscular mycorrhizae (endomycorrhizae)
Figure 37.13ba
Cortical cell
Epidermis
Cortex
Endodermis
Fungal vesicle
Fungal hyphae
Casparian strip
Root hair
10 m
Arbuscules
Plasma membrane
(LM)
Figure Mycorrhizae.
(b)
Arbuscular mycorrhizae (endomycorrhizae)

72. Cortical cell 10 m Arbuscules (LM) Figure 37.13bb
Figure Mycorrhizae.
Arbuscules
(LM)

73. Agricultural and Ecological Importance of Mycorrhizae
Farmers and foresters often inoculate seeds with fungal spores to promote formation of mycorrhizae
Some invasive exotic plants disrupt interactions between native plants and their mycorrhizal fungi
For example, garlic mustard slows growth of other plants by preventing the growth of mycorrhizal fungi
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74. Figure 37.14a
EXPERIMENT
Figure Inquiry: Does the invasive weed garlic mustard disrupt mutualistic associations between native tree seedlings and arbuscular mycorrhizal fungi?

75. Increase in plant biomass (%) Mycorrhizal colonization (%)
Figure 37.14b
RESULTS
300
200
Increase in plant biomass (%)
100
Invaded
Uninvaded
Sterilized invaded
Sterilized uninvaded
Soil type 40
Figure Inquiry: Does the invasive weed garlic mustard disrupt mutualistic associations between native tree seedlings and arbuscular mycorrhizal fungi? 30
Mycorrhizal colonization (%) 20
Seedlings 10
Sugar maple
Red maple
Invaded
Uninvaded
White ash
Soil type

76. 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
An epiphyte grows on another plant and obtains water and minerals from rain
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77. Staghorn fern, an epiphyte
Figure 37.15a
Figure Exploring: Unusual Nutritional Adaptations in Plants
Staghorn fern, an epiphyte

78. Parasitic plants absorb sugars and minerals from their living host plant
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79. Mistletoe, a photo- synthetic parasite
Figure 37.15b
Figure Exploring: Unusual Nutritional Adaptations in Plants
Mistletoe, a photo- synthetic parasite
Dodder, a nonphotosynthetic parasite (orange)
Indian pipe, a nonphoto- synthetic parasite of mycorrhizae

80. Mistletoe, a photo- synthetic parasite
Figure 37.15ba
Figure Exploring: Unusual Nutritional Adaptations in Plants
Mistletoe, a photo- synthetic parasite

81. Dodder, a nonphotosynthetic parasite (orange)
Figure 37.15bb
Figure Exploring: Unusual Nutritional Adaptations in Plants
Dodder, a nonphotosynthetic parasite (orange)

82. Indian pipe, a nonphoto- synthetic parasite of mycorrhizae
Figure 37.15bc
Figure Exploring: Unusual Nutritional Adaptations in Plants
Indian pipe, a nonphoto- synthetic parasite of mycorrhizae

83. Sundews Pitcher plants Venus flytrap Figure 37.15c
Figure Exploring: Unusual Nutritional Adaptations in Plants
Pitcher plants
Venus flytrap

84. Pitcher plants Figure 37.15ca
Figure Exploring: Unusual Nutritional Adaptations in Plants
Pitcher plants

85. Pitcher plants Figure 37.15cb
Figure Exploring: Unusual Nutritional Adaptations in Plants
Pitcher plants

86. Venus flytrap Figure 37.15cc
Figure Exploring: Unusual Nutritional Adaptations in Plants
Venus flytrap

87. Venus flytrap Figure 37.15cd
Figure Exploring: Unusual Nutritional Adaptations in Plants
Venus flytrap

88. Figure 37.15ce
Figure Exploring: Unusual Nutritional Adaptations in Plants
Sundews

89. Carnivorous plants are photosynthetic but obtain nitrogen by killing and digesting mostly insects
Video: Sun Dew Trapping Prey
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90. Denitrifying bacteria
Figure 37.UN01 N2
(from atmosphere)
(to atmosphere) N2
Nitrogen-fixing bacteria
H (from soil)
Denitrifying bacteria
NH4
NH3 (ammonia)
NH4 (ammonium)
NO3 (nitrate)
Ammonifying bacteria
Nitrifying bacteria
Figure 37.UN01 Summary figure, Concept 37.3
Organic material (humus)
Root

91. Figure 37.UN02
Figure 37.UN02 Appendix A: answer to Test Your Understanding, question 10