C h ap t e r 4 • The Terrestrial Environment    69

4 O
ur introduction of aquatic environments nutrients throughout the plant body. The topic of water balance
Chapter

was dominated by discussion of the physical and and the array of characteristics that plants and animals have
The Terrestrial Environment chemical properties of water—­characteristics such as
depth, flow rate, and salinity (see Chapter 3). When considering
evolved to overcome the problems of water loss are discussed
in more detail later (see Chapters 6 and 7).
the term terrestrial environment, however, people typically do Desiccation is not the only constraint imposed by the
not think of the physical and chemical characteristics of a place. transition from water to land. Because air is less dense than
What we most likely visualize is the vegetation: the tall, dense water, it results in a much lower drag (frictional resistance)
forests of the wet tropics; the changing colors of autumn in a on the movement of organisms; but it greatly increases the
temperate forest; or the broad expanses of grass that character- constraint imposed by gravitational forces. The upward force
ize the prairies. Animal life depends on the vegetation within a of buoyancy resulting from the displacement of water helps
region to provide the essential resources of food and cover—and organisms in aquatic environments overcome the constraints
as such, the structure and composition of plant life constrain imposed by gravity (see Section 3.2). In contrast, the need to
the distribution and abundance of animal life. But ultimately, as remain erect against gravitational force in terrestrial environ-
with aquatic environments, the physical and chemical features of ments results in a significant investment in structural materials
terrestrial environments set the constraints for life. Plant life is a such as skeletons (for animals) or cellulose (for plants). The
reflection of the climate and soils (as discussed in Chapter 6). giant kelp (Macrocystis pyrifera) inhabiting the waters off the
Regardless of the suitability of plant life for providing essential coast of California is an excellent example (Figure 4.1, left).
resources, the physical conditions within a region impose the It grows in dense stands called kelp forests. Anchored to the
primary constraints on animal life as well (Chapter 7). bottom sediments, these kelp (macroalgae) can grow 100 feet
We will explore key features of the terrestrial environ- or more toward the surface. The kelp are kept afloat by gas-
ment that directly influence life on land. Life emerged from filled bladders attached to each blade; yet when the kelp plants
the water to colonize the land more than a billion years ago. are removed from the water, they collapse into a mass. Lacking
The transition to terrestrial environments posed a unique set of supportive tissues strengthened by cellulose and lignin, the kelp
problems for organisms already adapted to an aquatic environ-
ment. To understand the constraints imposed by the terrestrial
environment, we must start by looking at the physical differ- Figure 4.1 (left) The giant kelp (Macrocystis pyrifera) inhabits
ences between the terrestrial and aquatic environments and at the waters off the coast of California. Anchored to the bottom
The input and decomposition of dead organic matter is a key factor in the development of forest soils. the problems these differences create for organisms making the sediments, these kelp plants can grow 100 feet or more toward
transition from water to land. the surface despite their lack of supportive tissues. These kelp
plants are kept afloat through the buoyancy of gas-filled bladders
attached to each blade, yet when the kelp plants are removed
4.1 Life on Land Imposes Unique from the water, they collapse into a mass. (right) In contrast, a
Chapter Guide Constraints
redwood tree (Sequoia sempervirens) of comparable height
allocates more than 80 percent of its biomass to supportive and
4.1 Life on Land Imposes Unique Constraints The transition from life in aquatic environments to life on land conductive tissues that help the tree resist gravitational forces.
brought with it a variety of constraints. Perhaps the greatest
4.2 Plant Cover Influences the Vertical Distribution of Light
constraint imposed by terrestrial environments is desiccation.
4.3 Soil Is the Foundation upon which All Terrestrial Life Depends Living cells, both plant and animal, contain about 75–95 percent
4.4 The Formation of Soil Begins with Weathering water. Unless the air is saturated with moisture, water readily
evaporates from the surfaces of cells via the process of diffusion
4.5 Soil Formation Involves Five Interrelated Factors (see Section 2.5). The water that is lost to the air must be re-
4.6 Soils Have Certain Distinguishing Physical Characteristics placed if the cell is to remain hydrated and continue to function.
Maintaining this balance of water between organisms and their
4.7 The Soil Body Has Horizontal Layers or Horizons
surrounding environment (referred to as an organism’s water
4.8 Moisture-Holding Capacity Is an Essential Feature of Soils balance) has been a major factor in the evolution of life on land.
4.9 Ion Exchange Capacity Is Important to Soil Fertility For example, in adapting to the terrestrial environment, plants
have evolved extensively specialized cells for different functions.
4.10 Basic Soil Formation Processes Produce Different Soils Aerial parts of most plants, such as stems and leaves, are coated
Ecological Issues & Applications Soil Errosion with a waxy cuticle that prevents water loss. While it reduces
water loss, the waxy surface also prevents gas exchange (carbon
dioxide and oxygen) from occurring. As a result, terrestrial plants
have evolved pores on the leaf surface (stomata) that allow gases
to diffuse from the air into the interior of the leaf (see Chapter 6).
To stay hydrated, an organism must replace water that
it has lost to the air. Terrestrial animals can acquire water by
drinking and eating. For plants, however, the process is passive.
Early in their evolution, land plants evolved vascular tissues
consisting of cells joined into tubes that transport water and
68

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cannot support its own weight under the forces of gravity. In absorption and reflection of solar radiation by plants. When 26 26 Figure 4.3 The concept of
contrast, a tree of equivalent height inhabiting the coastal for- walking into a forest in summer, you will observe a decrease 24 24
leaf area index (LAI). (a) A tree
est of California (Figure 4.1, right) must allocate more than in light (Figure 4.2a). You can observe much the same effect with a crown 10 m wide projects
22 22
80 percent of its total mass to supportive and conductive tissues if you examine the lowest layer in grassland or an old field a circle of the same size on the
20 20 ground. (b) Foliage density
in the trunk (bole), branches, leaves, and roots. (Figure 4.2b). The quantity and quality (spectral composition)
18 18 (area of leaves) at various
Another characteristic of terrestrial environments is their of light that does penetrate the canopy of vegetation to reach
16 16 heights above the ground.
high degree of variability, both in time and space. Temperature the ground varies with both the quantity and orientation of

Height (m)

Height (m)
14 14 (c) Contributions of layers in the
variations on land (air) are much greater than in water. The high the leaves. crown to the leaf area index.
12 12
specific heat of water prevents wide daily and seasonal fluctua- The amount of light at any depth in the canopy is affected (d) Calculation of leaf area
10 10
tions in the temperature of aquatic habitats (see Section 3.2). by the number of leaves above. As we move down through the index (LAI). The total leaf area is
8 8
In contrast, such fluctuations are a characteristic of air tem- canopy, the number of leaves above increases; so the amount 315 m2. The projected ground
peratures (see Chapter 2). Likewise, the timing and quantity of of light decreases. However, because leaves vary in size and 6 6
Total leaf area = 315 m2 area is 78.5 m2. The LAI is 4.
precipitation received at a location constrains the availability of shape, the number of leaves is not the best measure of quan- 4 4
water for terrestrial plants and animals as well as their ability tity. The quantity of leaves, or foliage density, is generally 2 2
to maintain water balance. These fluctuations in temperature expressed as the leaf area. Because most leaves are flat, the 0 0
25 50 75 100 0.25 0.5 0.75 1.00
and moisture have both a short-term effect on metabolic pro- leaf area is the surface area of one or both sides of the leaf. Leaf area (m2) Leaf area index
(b) (c)
cesses and a long-term influence on the evolution and distribu- When the leaves are not flat, the entire surface area is some-
tion of terrestrial plants and animals (see Chapters 6 and 7). times measured. To quantify the changes in light environment
Ultimately, the geographic variation in climate governs the with increasing area of leaves, we need to define the area of
large-scale distribution of plants and therefore the nature of ter- leaves per unit ground area (m2 leaf area/m2 ground area). This 10 m
total leaf area Leaf area index
restrial ecosystems (see Chapter 23). measure is the leaf area index ([LAI]; Figure 4.3). A LAI of projected ground area
=
(LAI)
3 indicates a quantity of 3 m2 of leaf area over each 1 m2 of
4.2 Plant Cover Influences the ground area. (d)

The greater the LAI above any surface, the lower the quan-
Vertical Distribution of Light tity of light reaching that surface. As you move from the top of
Ground area = πr 2 = 78.5 m2
(a)
In contrast to aquatic environments, where the absorption of the canopy to the ground in a forest, the cumulative leaf area
solar radiation by the water itself results in a distinct verti- and LAI increase. Correspondingly, light decreases. The gen-
cal gradient of light, the dominant factor influencing the eral relationship between available light and LAI is described
In addition to the quantity of light, the spectral composi- Length Length
vertical gradient of light in terrestrial environments is the by Beer’s law (see Quantifying Ecology 4.1).
tion (quality) of light varies through the plant canopy. Recall 0 0.5 1.0 0 0.5 1.0
that the wavelengths of approximately 400 to 700 nm make
10% reflected by
up visible light (Section 2.1 and Figure 2.1). These wave-
top of canopy 20% reflected by lengths are also known as photosynthetically active radiation Sunlight Sunlight
top of canopy (PAR) because they include the wavelengths used by plants
100% as a source of energy in photosynthesis (see Chapter 6).
The transmittance of PAR is typically less than 10 percent,
100%
whereas the transmittance of far-red radiation (730 nm) is
much greater. As a result, the ratio of red (660 nm) to far-red
5% radiation (R/FR ratio) decreases through the canopy. This
79%
shift in the spectral quality of light affects the production of
phytochrome (a pigment that allows a plant to perceive shad- Shade Shade
36%
ing by other plants), thus influencing patterns of growth and
allocation (see Chapter 6, Section 6.8). (a) Perpendicular (b) 60° angle
Besides the quantity of leaves, the orientation of leaves on
Figure 4.4 Influence of leaf orientation (angle) on the
the plant influences the attenuation of light through the canopy. interception of light energy. If a leaf that is perpendicular to the
The angle at which a leaf is oriented relative to the Sun changes source of light (a) intercepts 1.0 unit of light energy, the same leaf
7%
the amount of light it absorbs. If a leaf that is perpendicular to at an angle of 60 degrees relative to the light source will intercept
2% 34%
the Sun absorbs 1.0 unit of light energy (per unit leaf area/ only 0.5 unit (b). The reduction in intercepted light energy is a
time), the same leaf displayed at a 60-degree angle to the Sun result of the angled leaf projecting a smaller surface area relative
will absorb only 0.5 units. The reason is that the same leaf area to the light source.
(a) 2% (b) represents only half the projected surface area and therefore
5%
intercepts only half as much light energy (Figure 4.4). Thus,
Figure 4.2 Absorption and reflection of light by the plant canopy. (a) A mixed ­conifer–
leaf angle influences the vertical distribution of light through locations and at different times. For example, in high-latitude
deciduous forest reflects about 10 percent of the incident photosynthetically active radiation
(PAR) from the upper canopy, and it absorbs most of the remaining PAR within the canopy. the canopy as well as the total amount of light absorbed and environments, where sunlight angles are low, canopies having
(b) A meadow reflects 20 percent of the PAR from the upper surface. The middle and lower reflected. The sun angle varies, however, both geographically leaves that are displayed at an angle will absorb light more
regions, where the leaves are densest, absorb most of the rest. Only 2–5 percent of PAR (see Section 2.1) and through time at a given location (over effectively (see Figure 2.5). Leaves that are displayed at an
reaches the ground. the course of the day and seasonally). Consequently, different angle rather than perpendicular to the Sun are also typical of
(Adapted from Larcher 1980.) leaf angles are more effective at intercepting light in different arid tropical environments. In these hot and dry environments,

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Q uan ti fy i n g Ecol og y 4. 1 Beer’s Law and the Attenuation of Light

D ue to the absorption and reflection of light by leaves,

there is a distinct vertical gradient of light availability
from the top of a plant canopy to the ground. The greater the
1.0
24–26 m
24–26 m
Chlorophyll Particulates

0.8 (Top of canopy)

surface area of leaves, the less light will penetrate the canopy 15 kT = kw + kc + kd + kp

Net photosynthesis
Available light (ALi)

(μmol CO2 /m2/s)

and reach the ground. The vertical reduction, or attenuation, 22–24 m
of light through a stand of plants can be estimated using 0.6 22–24 m Total Water Dissolved
Beer’s law, which describes the attenuation of light through 10 20–22 m substance
a homogeneous medium. The medium in this case is the 0.4
20–22 m Ground level
canopy of leaves. Beer’s law can be applied to the problem of
5 18–20 m
light attenuation through a plant canopy using the following 18–20 m Whereas the light extinction coefficient for leaf area ex-
0.2
relationship: presses the attenuation of light per unit of LAI, these values
12–14 m 12–14 m of k are expressed as the attenuation of light per unit of water
0 1 2 3 4 0 0.2 0.4 0.6 0.8 1.0 depth (such as centimeter, meter, inches, or feet). Beer’s law
Leaf area index Leaf area index (LAIi) Available light (ALi) can then be used to estimate the quantity of light reaching any
above height i depth (z) by using the following equation:
Figure 1 Relationship between leaf area index above Figure 2 Relationship between available light (PAR) and
Light reaching any vertical rate of net photosynthesis at various heights in the canopy.
position i, expressed as a various heights in the canopy (LAIi) and the associated values aL z = e-ktz
ALi = e−LAIi × k of available light (ALi), expressed as a proportion of PAR at Available light is expressed as the proportion of PAR at the
proportion of light reaching the
top of the canopy the top of the canopy. top of the canopy (assumed to be 1500 μmol/m2/s). If the ecosystem supports submerged vegetation, such as
kelp (see Figure 4.1), seagrass, or other plants that are rooted in
Light extinction the bottom sediments, the preceding equation can be used to
coefficient calculate the available light at the top of the canopy. The equa-
tion describing the attenuation of light as a function of LAI can
For the example presented in Figure 4.3, we can construct downward, the amount of light reaching the leaves and the then be applied (combined) to calculate the further attenuation
The subscript i refers to the vertical height of the canopy. a curve describing the available light at any height in the can- corresponding rate of photosynthesis decline. from the top of the plant canopy to the sediment surface.
For example, if i were in units of meters, a value of i = 5 refers opy. In Figure 1, the light extinction coefficient has a value Beer’s law can also be used to describe the vertical at-
to a height of 5 m above the ground. The value e is the natural of k = 0.6 as an average value for a temperate deciduous for- tenuation of light in aquatic environments, but applying the
1. If we assume that the value of k used to calculate the verti-
logarithm (2.718). The light extinction coefficient, k, repre- est. We label vertical positions from the top of the canopy to light extinction coefficient (k) is more complex. The reduc-
cal profile of light in Figure 1 (k = 0.6) is for a plant canopy
sents the quantity of light attenuated per unit of leaf area ground level on the curve. Knowing the amount of leaves (LAI) tion of light with water depth is a function of various factors:
where the leaves are positioned horizontally (parallel to
index (LAI) and is a measure of the degree to which leaves above any position in the canopy (i), we can use the equation (1) attenuation by the water itself (see Section 3.3, Figure 3.7);
the forest floor), how would the value of k differ (higher
absorb and reflect light. The extinction coefficient will vary as to calculate the amount of light there. (2) attenuation by phytoplankton (microscopic plants sus-
or lower) for a forest where the leaves were oriented at a
a function of leaf angle (see Figure 4.4) and the optical prop- The availability of light at any point in the canopy will di- pended in water), typically expressed as the concentration
60-degree angle? (See the example in Figure 4.4.)
erties of the leaves. Although the value of ALi is expressed as rectly influence the levels of photosynthesis (see Figure 6.2). of chlorophyll (the light-harvesting pigment of plants) per
a proportion of the light reaching the top of the canopy, the The light levels and rates of light-limited photosynthesis volume of water (see Section 6.1); (3) attenuation by dissolved 2. In shallow-water ecosystems, storms and high wind can re-
quantity of light at any level can be calculated by multiplying for each of the vertical canopy positions are shown in the substances; and (4) attenuation by suspended particulates. sult in bottom sediments (particulates) being suspended in
this value by the actual quantity of light (or photosyntheti- curve in Figure 2. Light levels are expressed as a proportion Each of these factors has an associated light extinction coeffi- the water for some time before once again settling to the
cally active radiation) reaching the top of the canopy (units of of values for fully exposed leaves at the top of the canopy cient, and the overall light extinction coefficient (kT) is the sum bottom. How would this situation affect the value of kT and
μmol/m2/s). (1500 μmol/m2/s). As one moves from the top of the canopy of the individual coefficients: the attenuation of light in the water profile?

30 Jan Jul during the winter months. In these cases, the amount of light that
angled leaves reduce light interception during midday, when
Feb Aug penetrates a forest canopy varies with the season (Figure 4.6).
temperatures and demand for water are at their highest.
600 25 Mar Sep In early spring in temperate regions, when leaves are just ex-
Although light decreases downward through the plant
Apr Oct panding, 20–50 percent of the incoming light may reach the
canopy, some direct sunlight does penetrate openings in the
Photosynthetically active radiation

500 May Nov

Height above ground (m)

crown and reaches the ground as sunflecks. Sunflecks can ac- forest floor. In other regions characterized by distinct wet and
20 Jun Dec
400 count for 70–80 percent of solar energy reaching the ground in dry seasons, a similar pattern of increased light availability at the
forest environments (Figure 4.5). ground level occurs during the dry season (see Chapter 2).
(μmol/m2/s)

300 In many environments, seasonal changes strongly influence 15

leaf area. For example, in the temperate regions of the world, Figure 4.6 Measured vertical profiles of photosynthetically active
200
many forest tree species are deciduous, shedding their leaves 10 radiation (PAR) averaged monthly from 2003 to 2007 in Morgan-
100 Monroe State Forest (Indiana, United States). Values are expressed
as a proportion of PAR at the top of the canopy (27 m height). Note
0 Figure 4.5 Changes in the availability of light 5 that the profiles form two distinct groups corresponding to the
(photosynthetically active radiation [PAR]) at ground level in a presence or absence of foliage in this temperate deciduous forest.
lowland rain forest in Mexico over the course of a day. The spikes (Adapted from Wenze Yanga, Wenge Ni-Meistera, Nancy Y. Kiangb, Paul
6 8 10 12 14 16 18 20 0
result from sunflecks in an otherwise low-light. 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 R. Moorcroftc, Alan H. Strahler, Andrew Oliphante, Agricultural and forest
Time (hours) (Adapted from Chazdon and Pearcy 1991.) Proportion of available light (PAR) at top of canopy Meteorology Volume 150, Issues 7–8, 15 July 2010, Pages 895–907.)

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 74    Part o ne • T HE PHYSI CA L EN VI RONM ENT  C h ap t e r 4 • The Terrestrial Environment    75

4.3 Soil Is the Foundation upon Without appreciably influencing their composition, me-
chanical weathering breaks down rock and minerals into
Topography, the contour of the land, can affect how cli-
mate influences the weathering process. More water runs off
100

which All Terrestrial Life Depends smaller particles. Simultaneously, these particles are chemi- and less enters the soil on steep slopes than on level land; 90 10

Soil is the medium for plant growth; the principal factor con- cally altered and broken down through chemical weathering. whereas water draining from slopes enters the soil on low and 80 20
trolling the fate of water in terrestrial environments; nature’s The presence of water, oxygen, and acids resulting from the flat land. Steep slopes are also subject to soil erosion and soil
70 30
recycling system, which breaks down the waste products of activities of soil organisms and the continual addition of organic creep—the downslope movement of soil material that accumu- Clay

Pe
matter (dead plant and animal tissues) enhance the chemical lates on lower slopes and lowlands. 60 40

y
plants and animals and transforms them into their basic ele-

cla

rce
ments; and a habitat to a diversity of animal life, from small weathering process. Rainwater falling on and filtering through Time is a crucial element in soil formation: all of the fac-

nt
50

en
50 Silty
tors just listed assert themselves over time. The weathering of

s
this organic matter and mineral soil sets up a chain of chemical

rc
mammals to countless forms of microbial life (see Chapter 21).

ilt
Sandy clay

Pe
reactions that transform the composition of the original rocks rock material; the accumulation, decomposition, and mineral- 40 60
As familiar as it is, soil is hard to define. One definition clay
Silty clay
says that soil is a natural product formed and synthesized by the and minerals. ization of organic material; the loss of minerals from the upper 30 Sandy
Clay loam
loam 70
weathering of rocks and the action of living organisms. Another surface; and the downward movement of materials through clay loam
the soil all require considerable time. Forming well-developed 20 Loam 80
states that soil is a collection of natural bodies of earth, com- Sandy loam Silt loam
posed of mineral and organic matter and capable of supporting 4.5 Soil Formation Involves Five soils may require 2000 to 20,000 years. 10
Loamy Silt
90
plant growth. Indeed, one eminent soil scientist, Hans Jenny—a Sand sand
pioneer of modern soil studies—will not give an exact defini- Interrelated Factors 100 90 80 70 60 50 40 30 20 10
100

tion of soil. In his book The Soil Resource, he writes: Five interdependent factors are important in soil formation: 4.6 Soils Have Certain Percent sand

Popularly, soil is the stratum below the vegetation and

parent material, climate, biotic factors, topography, and time. Distinguishing Physical Figure 4.7 A soil texture chart showing the percentages of clay
Parent material is the material from which soil develops. The
above hard rock, but questions come quickly to mind. original parent material could originate from the underlying Characteristics (less than 0.002 mm), silt (0.002–0.05 mm), and sand (0.05–2.0 mm)
in the basic soil texture classes. For example, a soil with 60 percent
Many soils are bare of plants, temporarily or permanently, bedrock; from glacial deposits (till); from sand and silt car- Soils are distinguished by differences in their physical and sand, 30 percent silt, and 10 percent clay would be classified as a
or they may be at the bottom of a pond growing cattails. ried by the wind (eolian); from gravity moving material down chemical properties. Physical properties include color, texture, sandy loam.
Soil may be shallow or deep, but how deep? Soil may be a slope (colluvium); and from sediments carried by flowing structure, moisture, and depth. All may be highly variable from
stony, but surveyors (soil) exclude the larger stones. Most water (fluvial), including water in floodplains. The physical one soil to another. Interpreting Ecological Data
analyses pertain to fine earth only. Some pretend that soil character and chemical composition of the parent material are Color is one of the most easily defined and useful charac- Q1. What is the texture classification for a soil with 60 percent
in a flowerpot is not a soil, but soil material. It is embar- important in determining soil properties, especially during the teristics of soil. It has little direct influence on the function of a silt, 35 percent clay, and 5 percent sand?
rassing not to be able to agree on what soil is. In this, soil early stages of development. soil but can be used to relate chemical and physical properties. Q2. What is the texture classification for a soil with 60 percent
scientists are not alone. Biologists cannot agree on a defi- Biotic factors—plants, animals, bacteria, and fungi—all Organic matter (particularly humus) makes soil dark or black. clay and 40 percent silt?
nition of life and philosophers on philosophy. contribute to soil formation. Plant roots can function to break Other colors can indicate the chemical composition of the rocks
up parent material, enhancing the process of weathering, as and minerals from which the soil was formed. Oxides of iron
Of one fact we are sure. Soil is not just an abiotic environ-
well as stabilizing the soil surface and reducing erosion. Plant give a color to the soil ranging from yellowish-brown to red,
ment for plants. It is teeming with life—billions of minute and
roots pump nutrients up from soil depths and add them to the whereas manganese oxides give the soil a purplish to black color. Soil texture affects pore space in the soil, which plays a
not so minute animals, bacteria, and fungi. The interaction be-
surface. In doing so, plants recapture minerals carried deep Quartz, kaolin, gypsum, and carbonates of calcium and magne- major role in the movement of air and water in the soil and
tween the biotic and the abiotic makes the soil a living system.
into the soil by weathering processes. Through photosynthesis, sium give whitish and grayish colors to the soil. Blotches of vari- the penetration by roots. In an ideal soil, particles make up 50
Soil scientists recognize soil as a three-dimensional unit, or
plants capture the Sun’s energy and transfer some of this en- ous shades of yellowish-brown and gray indicate poorly drained percent of the soil’s total volume; the other 50 percent is pore
body, having length, width, and depth. In most places on Earth’s
ergy to the soil in the form of organic carbon. On the soil sur- soils or soils saturated by water. Soils are classified by color space. Pore space includes spaces within and between soil par-
surface, exposed rock has crumbled and broken down to produce
face, microorganisms break down the remains of dead plants using standardized color charts (i.e., Munsell soil color charts). ticles, as well as old root channels and animal burrows. Coarse-
a layer of unconsolidated debris overlaying the hard, unweath-
and animals that eventually become organic matter incorpo- Soil texture is the proportion of different-sized soil parti- textured soils have large pore spaces that favor rapid water
ered rock. This unconsolidated layer, called the regolith, varies
rated into the soil (see Chapter 21). Climate influences soil cles. Texture is partly inherited from parent material and partly infiltration, percolation, and drainage. To a point, the finer the
in depth from virtually nonexistent to tens of meters. This inter-
development both directly and indirectly. Temperature, precip- a result of the soil-forming process. Particles are classified on texture, the smaller the pores, and the greater the availability of
face between rock and the air, water, and living organisms that
itation, and winds directly influence the physical and chemical the basis of size into gravel, sand, silt, and clay. Gravel consists active surface for water adhesion and chemical activity. Very
characterizes the surface environment is where soil is formed.
reactions responsible for breaking down parent material and of particles larger than 2.0 mm, but they are not part of the fine fine-textured or heavy soils, such as clays, easily become com-
the subsequent leaching (movement of solutes through the fraction of soil. Soils are classified based on texture by defin- pacted if plowed, stirred, or walked on. They are poorly aerated
4.4 The Formation of Soil Begins soil) and movement of weathered materials. Water is essential ing the proportion of sand, silt, and clay. and difficult for roots to penetrate.
for the process of chemical weathering, and the greater the Sand ranges from 0.05 to 2.0 mm, is easy to see, and Soil depth varies across the landscape, depending on
with Weathering depth of water percolation, the greater the depth of weather- feels gritty. Silt consists of particles from 0.002 to 0.05 mm slope, weathering, parent materials, and vegetation. In grass-
Soil formation begins with the weathering of rocks and their ing and soil development. Temperature controls the rates of in diameter that can scarcely be seen by the naked eye; it feels lands, much of the organic matter added to the soil is from the
minerals. Weathering includes the mechanical destruction of rock biochemical reactions, affecting the balance between the accu- and looks like flour. Clay particles are less than 0.002 mm and deep, fibrous root systems of the grass plants. By contrast, tree
materials into smaller particles as well as their chemical modifi- mulation and breakdown of organic materials. Consequently, are too small to be seen under an ordinary microscope. Clay leaves falling on the forest floor are the principal source of
cation. Mechanical weathering results from the interaction of under conditions of warm temperatures and abundant water, controls the most important properties of soils, including its organic matter in forests. As a result, soils developed under na-
several forces. When exposed to the combined action of water, the processes of weathering, leaching, and plant growth (input water-holding capacity (see Section 4.8) and the exchange of tive grassland tend to be several meters deep, and soils devel-
wind, and temperature, rock surfaces flake and peel away. Water of organic matter) are maximized. In contrast, under cold, ions between soil particles and soil solution (see Section 4.9). oped under forests are shallow. On level ground at the bottom
seeps into crevices, freezes, expands, and cracks the rock into dry conditions, the influence of these processes is much more A soil’s texture is the percentage (by weight) of sand, silt, and of slopes and on alluvial plains, soils tend to be deep. Soils on
smaller pieces. Wind-borne particles, such as dust and sand, wear modest. Indirectly, climate influences a region’s plant and clay. Based on proportions of these components, soils are di- ridgetops and steep slopes tend to be shallow, with bedrock
away at the rock surface. Growing roots of trees split rock apart. animal life, both of which are important in soil development. vided into texture classes (Figure 4.7). close to the surface.

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4.7 The Soil Body Has Horizontal Below the organic layer is the A horizon, often referred
to as the topsoil. This is the first of the layers that are largely
retains much of the characteristics of the parent materials from
which it was formed. Below the C horizon lies the bedrock.
50
Field capacity
Layers or Horizons composed of mineral soil derived from the parent materi- 45

Initially, soil develops from undifferentiated parent material. als. In this horizon, organic matter (humus) leached from 40
4.8 Moisture-Holding Capacity Is

Water content (g/cm3)

Over time, changes occur from the surface down, through above accumulates in the mineral soil. The accumulation of 35
organic matter typically gives this horizon a darker color, Wilting point
the accumulation of organic matter near the surface and the
downward movement of material. These changes result in the distinguishing it from lower soil layers. Downward movement an Essential Feature of Soils 30
Available
water
formation of horizontal layers that are differentiated by physi- of water through this layer also results in the loss of miner- If you dig into the surface layer of a soil after a soaking rain, 25 capacity
cal, chemical, and biological characteristics. Collectively, a als and finer soil particles, such as clay, to lower portions of you should discover a sharp transition between wet surface soil
20
sequence of horizontal layers constitutes a soil profile. This the profile—­sometimes giving rise to an E horizon, a zone and the dry soil below. As rain falls on the surface, it moves into
pattern of horizontal layering, or horizons, is easily visible or layer of maximum leaching, or eluviation (from Latin ex the soil by infiltration. Water moves by gravity into the open 15
where a recent cut has been made along a road bank or during or e, “out,” and lavere, “to wash”) of minerals and finer soil pore spaces in the soil, and the size of the soil particles and 10
excavation for a building site (Figure 4.8). particles to lower portions of the profile. Such E horizons are their spacing determine how much water can flow in. Wide pore
quite common in soils developed under forests, but because of spacing at the soil surface increases the rate of water infiltration; 5
The simplest general representation of a soil profile con-
lower precipitation they rarely occur in soils developed under so coarse soils have a higher infiltration rate than fine soils do. Sand Loam Clay
sists of four horizons: O, A, B, and C (Figure 4.9). The surface
grasslands. If there is more water than the pore space can hold, we say Soil texture
layer is the O horizon, or organic layer. This horizon is domi- Coarse Fine
nated by organic material, consisting of partially decomposed Below the A (or E) horizon is the B horizon, also called that the soil is saturated, and excess water drains freely from
plant materials such as leaves, needles, twigs, mosses, and the subsoil. Containing less organic matter than the A horizon, the soil. If water fills all the pore spaces and is held there by Figure 4.10 Water content of three different soils at wilting
lichens. This horizon is often subdivided into a surface layer the B horizon shows accumulations of mineral particles such as internal capillary forces, the soil is at field capacity (physi- point (WP), field capacity (FC), and saturation. The three soils
clay and salts from the leaching from the topsoil. This process cally defined as the water content at –0.33 bar suction pressure, differ in texture from coarse-textured sand to fine-textured silty
composed of undecomposed leaves and twigs (Oi), a middle clay loam (see soil texture chart of Figure 4.7). Available water
layer composed of partially decomposed plant tissues (Oe), is called illuviation (from the Latin il, “in,” and lavere, “to or .0033 MPa). Field capacity is generally expressed as the
capacity (AWC) is defined as the difference between FC and WP.
and a bottom layer consisting of dark brown to black, homoge- wash”). The B horizon usually has a denser structure than the A percentage of the weight or volume of soil occupied by water
Both FC and WP increase from coarse- to fine-textured soils, and
neous organic material or the humus layer (Oa). This pattern of horizon, making it more difficult for plants to extend their roots when saturated compared to the oven-dried weight of the soil
the highest AWC is in the intermediate-textured soils.
layering is easily seen by carefully scraping away the surface downward. B horizons are distinguished on the basis of color, at a standard temperature. The amount of water a soil holds at
organic material on the forest floor. In temperate regions, the structure, and the kind of material that has accumulated as a field capacity varies with the soil’s texture—the proportion of Interpreting Ecological Data
organic layer is thickest in the fall, when new leaf litter ac- result of leaching from the horizons above. sand, silt, and clay. Coarse, sandy soil has large pores; water
Q1. Although fine-textured soils (silty clay loam) have a greater
cumulates on the surface. It is thinnest in the summer after The C horizon is the unconsolidated material that lies drains through it quickly. Clay soils have small pores and hold AWC, for this value to be achieved, the soil must be at or above
decomposition has taken place. under the subsoil and is generally made of original material considerably more water. Water held between soil particles by FC. In arid regions, low and infrequent precipitation may keep
from which the soil developed. Because it is below the zones capillary forces is capillary water. soil water content below FC for most of the growing season. If
of greatest biological activity and weathering and has not been As plants and evaporation from the soil surface extract the measured value of soil water content at a site is 10 g/cm3,
which soil texture (sand, silt, or clay) represented in Figure 4.10
sufficiently altered by the soil-forming processes, it typically capillary water, the amount of water in the soil declines. When
would have the greatest soil water available for uptake by plants?
the moisture level decreases to a point where plants can no
Q2. What if the value of soil water was 35 g/cm3?
longer extract water, the soil has reached the wilting point
(physically defined as the water content at –15 bar suction
Organic layer: dominated by pressure, or –1.5 MPa). The amount of water retained by the 4.9 Ion Exchange Capacity Is
organic material, consisting of soil between field capacity and wilting point (or the differ-
undecomposed or partially
ence between field capacity and wilting point) is the avail- Important to Soil Fertility
decomposed plant materials, O
such as dead leaves able water capacity (AWC), as shown in Figure 4.10. The Chemicals within the soil dissolve into the soil water to form a
A AWC provides an estimate of the water available for uptake by solution (see Section 3.5). Referred to as exchangeable nutri-
Topsoil: largely mineral soil
developed from parent material; plants. Although water still remains in the soil—filling up to 25 ents, these chemical nutrients in solution are the most readily
organic matter leached from percent of the pore spaces—soil particles hold it tightly, mak- available for uptake and use by plants (see Chapter 6). They are
above gives this horizon a ing it difficult to extract. held in soil by the simple attraction of oppositely charged par-
distinctive dark color B Both the field capacity and wilting point of a soil are heav- ticles and are constantly interchanging with the soil solution.
Subsoil: accumulation of ily influenced by soil texture. Particle size of the soil directly As described previously, an ion is a charged particle. Ions
mineral particles, such as clay influences the pore space and surface area onto which water carrying a positive charge are cations, and ions carrying a nega-
and salts leached from topsoil;
distinguished based on color, adheres. Sand has 30–40 percent of its volume in pore space, tive charge are anions. Chemical elements and compounds exist
structure, and kind of material whereas clays and loams (see soil texture chart in Figure 4.7) in the soil solution both as cations, such as calcium (Ca2+), mag-
accumulated from leaching range from 40 to 60 percent. As a result, fine-textured soils nesium (Mg2+), and ammonium (NH4+), and as anions, such as
C
Unconsolidated material have a higher field capacity than sandy soils, but the increased nitrate (NO3−) and sulfate (SO42−). The ability of these ions in
derived from the original surface area results in a higher value of the wilting point soil solution to bind to the surface of soil particles depends on
parent material from which the as well (see Figure 4.10). Conversely, coarse-textured soils the number of negatively or positively charged sites within the
soil developed
(sands) have a low field capacity and a low wilting point. Thus, soil. The total number of charged sites on soil particles within
Figure 4.9 A generalized soil profile. Over time, changes AWC is highest in intermediate clay loam soils. a volume of soil is called the ion exchange capacity. In most
Figure 4.8 The pattern of horizontal layering or soil horizons occur from the surface down, through the accumulation of organic The topographic position of a soil affects the movement of soils of the temperate zone, cation exchange predominates over
is easily visible where a recent cut has been made along a road matter near the surface and the downward movement of material. water both on and in the soil. Water tends to drain downslope, anion exchange because of the prevalence of negatively charged
bank. This soil is relatively shallow, with the parent material close These changes result in the formation of horizontal layers, or leaving soils on higher slopes and ridgetops relatively dry and particles in the soil, referred to as colloids. The total number of
to the surface. horizons. creating a moisture gradient from ridgetops to streams. negatively charged sites, located on the leading edges of clay

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particles and soil organic matter (humus particles), is called the is one of the most familiar of all chemical conditions in the Entisol Aridisol Vertisol
cation exchange capacity (CEC). These negative charges en- soil. Typically, soils range from pH 3 (extremely acid) to pH 9
able a soil to prevent the leaching of its positively charged nutri- (strongly alkaline). Soils of more than pH 7 (neutral) are consid- Immature soils that lack Develop in very dry Dark clay soils that show
vertical development of environments; low in significant expansion and
ent cations. Because in most soils there are far fewer positively ered basic, and those of pH 5.6 or less are acid. As soil acidity horizons; associated organic matter; high contraction due to wetting
charged than negatively charged sites, anions such as nitrate increases, the proportion of exchangeable Al3+ increases, and with recently deposited in base content; prone and drying
(NO3−) and phosphate (PO34−) are not retained on exchange Ca2+, Na+, and other cations decrease. High aluminum (Al3+) sediments to the process of
salinization
sites in soils but tend to leach away quickly if not taken up by concentrations in soil solution can be toxic to plants. Aluminum
plants. The CEC is a basic measure of soil quality and increases toxicity damages the root system first, making the roots short,
with higher clay and organic matter content. thick, and stubby. The result is reduced nutrient uptake.
Mollisol Inceptisol Spodosol
Cations occupying the negatively charged particles in the
soil are in a state of dynamic equilibrium with similar cations Surface horizons dark Young soils that are Light gray, whitish surface
in the soil solution (Figure 4.11). Cations in soil solution are 4.10 Basic Soil Formation brown to black with soft
consistency; rich in
more developed than
entisols; often shallow;
horizon on top of black
or reddish B horizon;
continuously being replaced by or exchanged with cations
on the clay and humus particles. The relative abundance of Processes Produce Different Soils bases; soils of semi-humid
regions; prone to the
moderate development
of horizons
high in extractable iron
and aluminum; formed
different ions on exchange sites is a function of their con- process of calcification through the process of
Broad regional differences in geology, climate, and vegetation podzolization
centration in the soil solution and the relative affinity of each give rise to characteristically different soils. The broadest level
ion for the sites. In general, the physically smaller the ion of soil classification is the order. Each order has distinctive
and the greater its positive charge, the more tightly it is held. features, summarized in Figure 4.12, and its own distribution, Alfisol Histosol Ultisol
The lyotropic series places the major cations in order of their mapped in Figure 4.13. Although a wide variety of processes Shallow penetration of High content of organic Intensely leached; strong
strength of bonding to the cation exchange sites in the soil: are involved in soil formation (pedogenesis), soil scientists humus; translocation matter; formed in areas clay translocation; low
recognize five main soil-forming processes that give rise to of clay; well-developed with poor drainage; base content; humid warm
Al3 + 7 H + 7 Ca2 + 7 Mg 2 + 7 K + = NH4+ 7 Na+ horizons bog and muck soils climate; formed by
these different classes of soils. These processes are laterization, process of laterization
However, higher concentrations in the soil solution can over- calcification, salinization, podzolization, and gleization.
come these differences in affinity. Laterization is a process common to soils found in humid
Hydrogen ions added by rainwater, by acids from organic environments in the tropical and subtropical regions. The hot,
matter, and by metabolic acids from roots and microorganisms rainy conditions cause rapid weathering of rocks and minerals. Andisol Oxisol Gelisol
increase the concentration of hydrogen ions in the soil solution Movements of large amounts of water through the soil cause
Developed from volcanic Highly weathered soils Presence of permafrost
and displace other cations, such as Ca2+, on the soil exchange heavy leaching, and most of the compounds and nutrients made parent material; not with nearly featureless or soil temperature of 0°C
sites. As more and more hydrogen ions replace other cations, available by the weathering process are transported out of the highly weathered; upper profile; red, yellow, or or less within 2 meters
the soil becomes increasingly acidic (see Section 3.7). Acidity soil profile if not taken up by plants. The two exceptions to this layers dark colored; gray; rich in kalolinate, of the surface; formed
low bulk density iron oxides, and often through the process
humus; in tropics and of gleization
subtropics

Figure 4.12 Profiles and general description of the 12 major soil orders of the world.
− K+
Soil particle
− Ca2+
− Figure 4.13 The world
− − − distribution of the 12 major soil
− −
− Mg2+ orders shown in Figure 4.12.
Air space K+ Ca2+
K+ H+ (Adapted from USGS, Soil
Conservation Service.)
Water available
to plant H2O + CO2 H2CO3 HCO3− + H+

Root hair

Soil particle
surrounded Root hair
by film of water
Cell wall

(a) Water film around soil particles (b) Cation exchange in soil

Figure 4.11 The process of cation exchange in soils. Cations occupying the negatively Alfisols Histosols Oxisols Vertisols Rocky soil
charged particles in the soil are in a state of dynamic equilibrium with similar cations in the
soil solution. Cations in soil solution are continuously being replaced by or exchanged with Aridisols Inceptisols Spodosols Gelisols Icefields/glaciers
cations on clay and humus particles. Cations in the soil solution are also taken up by plants Entisols Mollisols Ultisols Andisols Shifting sands
and leached to ground and surface waters.

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process are compounds of iron and aluminum. Iron oxides give By 1935 an additional 850 million tons of topsoil were re- Under this federal project, nearly 220 million trees were
tropical soils their unique reddish coloring (see Ultisol profile moved by wind erosion. It is estimated that by 1935 wind ero- planted, creating more than 18,000 miles of windbreaks on
in Figure 4.12). Heavy leaching also causes these soils to be sion had damaged 66 million hectares across 80 percent of the some 30,000 farms, which formed a “shelter belt” from the
acidic because of the loss of other cations (other than H1). High Plains. By 1938 it was estimated that 12.5 inches of top- Texas Panhandle to the Canadian border.
Calcification occurs when evaporation and water uptake soil had been lost over an area of 4 million hectares and 6.5 cm Although the end of the drought, together with soil con-
by plants exceed precipitation. The net result is an upward had been lost over another 5.5 million hectares. servation efforts following the Dust Bowl, abated the dramatic
movement of dissolved alkaline salts, typically calcium carbon- The storms generated by this environmental disaster dark- dust storms that blackened the skies over North America, the
ate (CaCO3), from the groundwater. At the same time, the infil- ened cities, buried homes and farm equipment, killed live- problem of soil erosion on agricultural lands remains a seri-
tration of water from the surface causes a downward movement stock, and represented a serious health risk (Figure 4.15b ous environmental issue. Approximately 50 percent of Earth’s
of the salts. The net result is the deposition and buildup of these and c). Overall, the Dust Bowl rendered millions of acres land surface is devoted to agriculture, with about one-third
deposits in the B horizon (subsoil). In some cases, these depos- of farmland virtually useless, left roughly half a million planted in crops and two-thirds used for grazing. Of these
its can form a hard layer called caliche (Figure 4.14 top). Americans homeless, and forced hundreds of thousands of two areas, cropland is more susceptible to erosion because the
Salinization is a process that functions similar to calcifi- people off the land. It also resulted in the most intense period vegetation is most often removed and the soil tilled (plowed)
cation, only in much drier climates. It differs from calcifica- of internal migration in U.S. history. Between 1932 and 1940, before crops are planted. This functions to destabilize the soil
tion in that the salt deposits occur at or near the soil surface it is estimated that 2.5 million people abandoned the plains for surface, increasing rates of erosion resulting from both wind
(Figure 4.14 bottom). Saline soils are common in deserts other regions of the country. and water (Figure 4.16a). In addition, croplands are often
but may also occur in coastal regions as a result of sea spray. In response to the environmental disaster of the Dust left without vegetation cover between plantings (exposing the
Salinization is also a growing problem in agricultural areas Bowl, U.S. president, Franklin Delano Roosevelt, established bare soil surface to erosion). According to David Pimentel of
where irrigation is practiced. the Soil Erosion Service (later the Soil Conservation Service, Cornell University, one of the leading experts in the study of
Podzolization occurs in cool, moist climates of the mid- and now the Natural Resources Conservation Service), which agricultural ecology, currently about 80 percent of the world’s
latitude regions where coniferous vegetation (e.g., pine forests) marked the first major federal commitment to the preservation agricultural land suffers moderate to severe soil erosion.
dominates. The organic matter of coniferous vegetation creates of natural resources in private hands. Even more significantly, Worldwide, erosion on cropland averages about 30 tons per
strongly acidic conditions. The acidic soil solution enhances the Figure 4.14 (top) In arid regions, salinization occurs when salts in 1935, the Prairie States Forestry Project was established. hectare per year and ranges from 0.5 to 400 tons per hectare
process of leaching, causing the removal of cations and com- (the white crust at the center of the photo) accumulate near the
pounds of iron and aluminum from the A horizon (topsoil). This soil surface because of surface evaporation. (bottom) Calcification
process creates a sublayer in the A horizon that is composed of occurs when calcium carbonates precipitate out from water
white- to gray-colored sand (see Spodosol profile in Figure 4.12). moving downward through the soil or from capillary water moving
upward from below. The result is an accumulation of calcium in
Gleization occurs in regions with high rainfall or low-
the B horizon (seen as the white soil layer in the photo).
lying areas associated with poor drainage (waterlogged). The Wash.
constantly wet conditions slow the breakdown of organic mat- Montana
North
ter by decomposers (bacteria and fungi), allowing the matter These five processes represent the integration of climate Dakota
Oregon
to accumulate in upper layers of the soil. The accumulated and edaphic (relating to the soil) factors on the formation South
Idaho
Wyoming Dakota
organic matter releases organic acids that react with iron in the of soils, giving rise to the geographic diversity of soils that
soil, giving the soil a black to bluish-gray color (see Gelisol influence the distribution, abundance, and productivity of ter- Nevada Nebraska
profile in Figure 4.12 as an example of soil formed through the restrial ecosystems. (We will explore these topics further in Utah Colorado
process of gleization). Chapters 20, 21, and 23.) Calif. Kansas

New Okla. (b)

Arizona
Mexico

Texas
Eco l o g i c a l Soil Erosion Is a Threat to Agricultural Sustainability
Issues & Applications 0 400 miles

In a report released in 1909, the U.S. Bureau of Soils stated native grassland was destroyed, the severe drought conditions
“The soil is the one indestructible, immutable asset that the na- and high winds during the period of the 1930s resulted in an in-
tion possesses. It is the one resource that cannot be exhausted; creased susceptibility of the topsoil to wind erosion. As a result, The Dust Bowl 1930–1939
that cannot be used up.” Yet less than three decades later, the dust storms raged nearly everywhere across the Great Plains Most affected area
loss of soil resources would be at the center of one of the worst of North America; but the most severely affected areas were in
Other affected area
environmental disasters in U.S. history—the Dust Bowl; a the Oklahoma and Texas panhandles, western Kansas, eastern
disaster that would have profound economic, social, and envi- Colorado, and northeastern New Mexico—a region that would
(a) (c)
ronmental costs. become known as the Dust Bowl (Figure 4.15a). The most
Between 1909 and 1929 farmers had tilled some 13 ­million severe dust storms occurred between 1935 and 1938, although Figure 4.15 (a) Map of the region of the Great Plains known as the Dust Bowl. This
hectares of land in the Great Plains. In doing so they destroyed they would continue through 1941. It was estimated that 300 represented the agricultural region most affected by soil erosion during the drought years
the sod—the grass-covered surface soil held together by the million tons of soil were removed from the region in May 1934 from 1930 to 1939. Photographs of (b) dust storms (Boise City, Oklahoma, on April 15, 1935,
dense mat of fibrous roots. Once this protective cover of the and spread over large portions of the eastern United States. and (c) their aftermath (Guymon, Oklahoma, March 29, 1937).

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per year. As a result of soil erosion, during the past four de- the susceptibility of the soil surface to erosion. Soil structure
cades about 30 percent of the world’s arable land has become influences the ease with which soils can be eroded. Soils with S u m m a ry
unproductive, much of which has been abandoned for agricul- medium-to-fine texture (see Section 4.6) and low organic
tural use. Each year an estimated 10 million hectares of crop- matter content are most easily eroded. Typically these soils Life on Land 4.1 it can be used to relate chemical and physical properties. Soil
land worldwide are abandoned because of lack of productivity have low water infiltration rates and are therefore susceptible Maintaining the balance of water between organisms and their texture is the proportion of different-sized soil particles—sand,
caused by soil erosion. to high rates of erosion by water and displacement by wind. surrounding environment has been a major influence on the silt, and clay. A soil’s texture is largely determined by the par-
Rates of soil erosion on agricultural lands are influenced Plant cover, both living and dead, greatly reduces rates of evolution of life on land. The need to remain erect against ent material but is also influenced by the soil-forming process.
by a variety of factors. Topography of the landscape, patterns erosion by protecting the soil surface from exposure to agents the force of gravity in terrestrial environments results in a Soil depth varies across the landscape, depending on slope,
of rainfall and wind, and exposure all combine to influence of erosion. significant investment in structural materials. Variations in weathering, parent materials, and vegetation.
Current estimates suggest that the degradation of agri- temperature and precipitation have both a short-term effect on
cultural lands alone will depress world food production by Soil Horizons 4.7
metabolic processes and a long-term influence on the evolution
Figure 4.16 (a) An agricultural field that has been recently approximately 30 percent over the next 50 years, while during and distribution of terrestrial plants and animals. The result is Soils develop in layers called horizons. Four horizons are
tilled. As a result of the lack of cover, tilled fields have high rates that same period the world population is predicted to exceed a distinct pattern of terrestrial ecosystems across geographic commonly recognized, although not all of them are necessar-
of erosion from wind and water. Some methods being adopted 9 billion (United Nations medium scenario; see Chapter 11, gradients of temperature and precipitation. ily present in any one soil: the O or organic layer; the A (some-
in the effort to reduce soil erosion and promote sustainable Ecological Issues & Applications). These forecasts point to times E) horizon, or topsoil, characterized by accumulation
agricultural practices are (b) contour and (c) no-till farming. the need to develop soil conservation techniques known to of organic matter; the B horizon, or subsoil, in which mineral
Light 4.2
dramatically reduce soil erosion. For example, commercial Light passing through a canopy of vegetation becomes attenu- materials accumulate; and the C horizon, the unconsolidated
corn production in the United States, which uses a practice of ated. The density and orientation of leaves in a plant canopy material underlying the subsoil and extending downward to
continuous crop production with annual plowing and removal influence the amount of light reaching the ground. Foliage den- the bedrock.
of all plant materials at harvest, results in an average soil ero- sity is expressed as leaf area index (LAI), the area of leaves per
sion rate of 44 tons per hectare per year. By using a practice Moisture-Holding Capacity 4.8
unit of ground area. The amount of light reaching the ground
of crop rotation in which a series of dissimilar/different types in terrestrial vegetation varies with the season. In forests, only The amount of water a soil can hold is one of its important
of crops are planted in the same area in sequential seasons about 1–5 percent of light striking the canopy reaches the characteristics. When water fills all pore spaces, the soil is
(e.g., corn, wheat, and hay) erosion rates have been shown to ground. Sunflecks on the forest floor enable plants to endure saturated. When a soil holds the maximum amount of water it
decline to as little as 3 tons per hectare per year. No-till tech- shaded conditions. can retain, it is at field capacity. Water held between soil par-
niques, in which crops are planted directly in the soil without ticles by capillary forces is capillary water. When the moisture
tilling or plowing the ground (Figure 4.16b), reduce average Soil Defined 4.3 level is at a point where plants cannot extract water, the soil has
rates of erosion to 0.14 tons per hectare per year in corn fields. reached wilting point. The amount of water retained between
Soil is a natural product of unconsolidated mineral and organic
Similar reductions in rates of erosion have been measured with field capacity and wilting point is the available water capacity.
matter on Earth’s surface. It is the medium for plant growth; the
contour planting (plowing and planting row crops on a contour The available water capacity of a soil is a function of its texture.
principal factor controlling the fate of water in terrestrial envi-
rather than up and down hill; Figure 4.16c) and the use of
ronments; nature’s recycling system, which breaks down the Ion Exchange 4.9
grass strips between crop rows. What all of these techniques
waste products of plants and animals and transforms them into
share in common is that they serve to protect the soil surface Soil particles, particularly clay particles and organic matter,
their basic elements; and a habitat to a diversity of animal life.
from direct exposure to wind and rain. are important to nutrient availability and the cation exchange
capacity of the soil—the number of negatively charged sites on
(a) Weathering 4.4
soil particles that can attract positively charged ions. Cations
Soil formation begins with the weathering of rock and miner- occupying the negatively charged particles in the soil are in a
als. In mechanical weathering, water, wind, temperature, and state of dynamic equilibrium with similar cations in the soil
plants break down rock. In chemical weathering, the activity solution. Percent base saturation is the percentage of sites oc-
of soil organisms, the acids they produce, and rainwater break cupied by ions other than hydrogen.
down primary minerals.
Soil Formation Processes Form Different Soils 4.10
Soil Formation 4.5 Broad regional differences in geology, climate, and vegetation
Soil results from the interaction of five factors: parent material, give rise to characteristically different soils. The broadest level
climate, biotic factors, topography, and time. Parent mate- of soil classification is the order. Each order has distinctive
rial provides the substrate from which soil develops. Climate features. Soil scientists recognize five main soil-forming pro-
shapes soil development through temperature, precipitation, cesses that give rise to these different classes of soils. These
and its influence on vegetation and animal life. Biotic factors— processes are laterization, calcification, salinization, podzoliza-
vegetation, animals, bacteria, and fungi—add organic matter tion, and gleization.
and mix it with mineral matter. Topography influences the
amount of water entering the soil and the rates of erosion. Time Soil Erosion Ecological Issues & Applications
is required to fully develop distinctive soils. Soil erosion on agricultural lands is a serious environmental
problem. The removal of natural vegetation and the plowing
Distinguishing Characteristics 4.6 of the soil destabilizes the soil surface and greatly enhances
Soils differ in the physical properties of color, texture, and erosion from wind and water. Sustainable practices such as
(b) (c)
depth. Although color has little direct influence on soil function, contour and no-till farming can greatly reduce rates of soil loss.

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 84    Part o ne • T HE PHYSI CA L EN VI RONM ENT

Study Questions
1. How do differences in the characteristics of terrestrial and 6. Use Figure 4.10 to answer this question: Which soil holds
aquatic environments influence the form and function of more moisture at field capacity: clay or sand? Which
organisms? soil holds more moisture at wilting point: clay or sand?
2. Assume that two forests have the same quantity of Which soil type has a greater availability of water for
leaves (leaf area index). In one forest, however, the plant ­uptake when the water content of the soil is 3.0 in/ft
leaves are oriented horizontally (parallel to the forest (value on y-axis)?
floor). In the other forest, the leaves are positioned at 7. Describe the E horizon of soil, and state the environments
an angle of 60 degrees. How would the availability of where we might expect to observe this.
light at the forest floor differ for these two forests at 8. Describe the terms field capacity, capillary water, wilting
noon? In which forest would the leaves at the bottom point, and available water capacity.
of the canopy (lower in the tree) receive more light at 9. How does pH influence the base saturation of a soil?
mid-morning? 10. Why is the process of salinization more prevalent in arid
3. What is the general shape of the curve that describes the areas? How does irrigation increase the process of salini-
vertical attenuation of light through the plant canopy zation in agricultural areas?
based on Beer’s law? Why is it not a straight line (linear)? 11. What soil-forming process is dominant in the wet tropical
4. Define soil. regions? How does this process influence the availability
5. What role does weathering play in soil formation? What of nutrients to plant roots in the A horizon?
factors are involved in the process of weathering?

Further Readings
Classic Studies Kohnke, H., and D. P. Franzmeier. 1994. Soil science
Wilde, S. A. 1946. Forest Soils and Forest Growth. Waltham, ­simplified. Prospect Heights, IL: Wavelength Press.
MA: Chronica Botanica. A well-written and illustrated overview of concepts and
A classic text on soils that was influential on the development of ­principles of soil science for the general reader. Provides many
soil science. examples and applications of basic concepts.
Patton, T. R. 1996. Soils: A new global view. New Haven, CT:
Current Research Yale University Press.
Brady, N. C., and R. W. Weil. 2008. The nature and properties Presents a new view and approach to studying soil formation at
of soils. 14th ed. Upper Saddle River, NJ: Prentice Hall. a global scale.
The classic introductory textbook on soils. Used for courses in
soil science.
Jenny, H. 1994. Factors of soil formation. Mineola, NY: Dover
Publications.
A well-written and accessible book by one of the pioneers in soil
science.

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