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MODULE 5

SOIL ORGANISM AND ORGANIC MATTER

Intended Learning Objectives

At the end of the chapter the students will be able to:


1. Identify and describe soil organism and Organic Matter
2. Assess and examine the soil organism and Organic Matter
3. Evaluate and characterize the soil organism and organic matter

Introduction

The Soil Organic Matter and Soil Biology

Soil is teeming with life, both macroscopic and microscopic. These life forms range in size
from invisible microorganisms to easily visible insects, earthworms and plant roots. In a teaspoon
of soil, there are millions of bacteria, hundreds of thousands of fungi, thousands of protozoa and
many larger organisms. These soil organisms play essential roles in nutrient cycling and energy
flow, both of which influence soil fertility and crop production.

Soil organic matter and soil organisms are inextricably connected. Microbial biomass is
the living component of soil organic matter, and microorganisms are the catalysts for most
nutrient-releasing processes. They make it possible for crops to grow and for soils to be
productive. On the other hand, microbial growth and activities depend on available carbon and
other mineral nutrients as well as a favorable physical and chemical environment. The way soil is
managed through tillage and cropping systems has a profound impact on life below ground.

Soil Organic Matter as Food

Soil organisms form a complex food web, and soil organic matter is the base of the web.
Most soil microorganisms use organic compounds in soil organic matter as carbon and energy
sources. Some soil organisms feed directly on living roots, but most depend on dead plant matter.
Small insects such as the springtail, a micro-invertebrate, break up plant residue into small pieces,
which accelerates further decomposition by microorganisms. Within the soil food web, there are
also carnivores, parasites and predators. As in an aboveground ecosystem, these organisms are
interdependent and help cycle nutrients from organic to inorganic forms that are available to
crops.

Except for the area next to the root, called the rhizosphere, soil is a nutrient-poor
environment for microbial growth. Nutrients and carbon, in the form of plant and animal residues,
tend to enter the soil intermittently. Consequently, microorganisms are faced with a feast-or-
famine existence. Soil microorganisms respond rapidly to the addition of plant and animal
residues. They break down complex organic compounds, such as cellulose and lignin in plant
residues, into simple organic compounds. Some of the carbon in these simple organic compounds
becomes part of the microbial biomass and provides energy for microbial growth. Some becomes
carbon dioxide.

Soil Science I: 1
The more stable fraction of soil organic matter, humus, is also a source of carbon for
microorganisms. Organic compounds found in humus have complex chemical structures and are
more resistant to decomposition than fresh plant or animal residues. Humus is also associated
with mineral particles and forms materials called humate-clay complexes that protect the organic
matter from decomposition by soil microorganisms. Therefore, humus serves as a slow-release
source of carbon and energy.

In addition to carbon, soil organic matter contains substantial amounts of organic nitrogen,
phosphorus, sulfur and many trace elements. Microorganisms perform an important function in
cycling these nutrients. They convert organically bound elements to inorganic or mineral forms
that are available for plant use. This process is called mineralization. Microorganisms, as well as
plants, also immobilize nutrients in their biomass as they grow. These nutrients are unavailable to
plants until the microorganisms and plants die and decompose. Mineralization and immobilization
are key processes in nutrient cycling.

Nitrogen is the nutrient in highest demand by plants. Plants mainly use inorganic forms of
nitrogen, such as ammonium and nitrate, which are products of microbial transformations. Plants
and microbes use ammonium from nitrogen fixation and mineralization to form proteins, nucleic
acids and cell walls. Nitrifying bacteria convert ammonium to nitrate in a process called
nitrification. Microorganisms also convert nitrate to various gases, N2, NO, N2O, through a
process called denitrification. This process occurs when soils are water saturated and oxygen is
low. Denitrification causes nitrogen losses from the soil to the atmosphere.

Microorganisms compete with plants for nitrogen. During the decomposition of organic
residues, microbial needs for nitrogen are met first. This is why with low-nitrogen residues, most
nitrogen is immobilized and not available for plant use. Nitrogen not used by microbes is released
to the soil and becomes available for plant use.

The increase in soil organic matter that occurs in conservation tillage systems results in
greater soil biological activity and soil biodiversity. Generally speaking, microbial biomass
increases along with soil organic matter and makes up 1–4 percent of the total organic matter.
Reducing tillage increases the amount of microbial biomass in soil. This improves soil quality and
promotes a constant cycling of nutrients, some of which are available for crop growth.

On the basis of organic matter content, soils are characterized as mineral or organic.
Mineral soils form most of the world’s cultivated land and may contain from a trace to 30 percent
organic matter. Organic soils are naturally rich in organic matter principally for climatic reasons.
Although they contain more than 30 percent organic matter, it is precisely for this reason that they
are not vital cropping soils. This soils bulletin concentrates on the organic matter dynamics of
cropping soils. In brief, it discusses circumstances that deplete organic matter and the negative
outcomes of this. The bulletin then moves on to more proactive solutions. It reviews a “basket” of
practices in order to show how they can increase organic matter content and discusses the land
and cropping benefits that then accrue.

Soil Science I: 2
LESSON I
SOIL ORGANISMS

The soil is a complex system of organic and inorganic matter. It consists of various layers
of this material, each varying in the amount of solid, gases, liquids, and organic matter. A general
analysis of soil shows that about 40% are rocks and minerals, 25% gases, 25% liquid, and 10%
organic matter. This combination creates the soil environment, which has several characteristics
that make it worthwhile to study. It provides a place for plants to secure themselves. Soil acts as a
filter that helps purify the water, and it is the place where many nutrients are recycled. Finally the
topic of this paper, soil is the habitat for thousands of different organisms around the world.
➢ Soil Organisms are generally grouped into two categories:
1. micro fauna/microorganisms and
2. macrofauna

➢ The main soil microorganisms include

a. bacteria,
b. fungi, and
c. protozoa.

➢ The macro fauna include:


a. oligochaeta,
b. arthropods,
c. mollusks, and
d. nematods.
There are many other organisms that spend some time in the soil, but usually just for
reproduction or feeding, and are not included in this paper. The organisms mentioned above play
an essential role in soil formation and the soil environment. The larger organisms tend to depend
more on the microorganisms that are part of the lower trophic levels. Therefore, microorganisms
will be discussed first to provide background information.

Soil Science I: 3
Perhaps the most important microorganisms in the soil ecosystem are bacteria. They are
single-celled organisms that are responsible for many important processes. One thing they do is
decompose plant material as well as other organisms' waste. These waste materials contain
nutrients, and the bacteria's cells are able to convert them from an unusable form into a form that
can be used. One essential nutrient that is converted by bacteria is nitrogen. Nitrogen-fixing
bacteria are able to use the nitrogen that exists in the atmosphere (N2) and change it through
cellular processes into ammonia (NH3). Nitrifying bacteria mix ammonia (NH3) with water
(H2O) to create ammonium ion (NH4+) and nitrate (NO3-), which can be used by other
organisms. Protozoa are also present in the soil environment and contribute to distributing
ammonium into the soil for use by plants and other organisms. They feed on bacteria but do not
need as much nitrogen as the bacteria provide, and therefore release the excess ammonium. Plants
need this to grow, so much so, that humans began putting more nitrogen into fertilizers through an
industrial process created by Fritz Haber and Carl Bosch. Denitrifying bacteria are responsible for
putting the nitrogen back into the atmosphere as N2.

Bacteria are also responsible for decomposing other compounds as well. One phylum of
bacteria called actinobacteria or actinomycetes are responsible for decomposing chitin, cellulose,
and other difficult materials in dead plants, animals and fungi.

Other organisms cannot disintegrate these materials because of their tough structure. The
bacteria are able to produce an enzyme allowing them to reduce the chitin, for example, from a
polymer into simple sugars and ammonia that other organisms can then use. Actinomycetes are a
type of bacteria that used to be considered as fungi. They were thought to be so because they grow
hyphae like fungi, and they can live in harsh environments in which other bacteria usually do not
survive, like higher pH levels and dryer conditions. They since were reclassified and renamed
actinobacteria. These bacteria give the soil its earthy scent, which actually indicates a healthy
environment. Bacteria such as this help to maintain healthy soil environments for plants, allowing
them to get the nitrogen they need, and decomposing organic waste.
Fungi serve in a similar capacity in the soil environment. They tend to be more abundant
in places where the bacteria cannot thrive, such as environments that are acidic, have low amounts
of nitrogen, have low moisture, and high complex carbohydrates (bacteria cannot break them
down). Fungi fill the niche of decomposing tough organic matter such as chitin, cellulose, keratin
and lignins. Again, most organisms cannot digest these materials, but fungi, like actinobacteria,
can produce enzymes that allow them to degrade those strong compounds into sugars that can be
digested. Not only are they decomposers, but fungi also are responsible for supplying nutrients to
other organisms, in this case plants. Fungi form a mutualistic relationship with plants called
mycorriza, which allows both parties to benefit. The fungi grow on or within (endopyhtic) the
roots of the plant. Because of the hyphae's high surface area to volume ratio, fungi are able to
better access essential nutrients and deliver them to the plants. The plants, in turn, give the fungi
food in the form of glucose.
Already, the importance of these microorganisms is clear, but in addition to them
converting nutrients into usable forms, they also are the food source for many of the macro fauna
in the soil. The macro fauna depends on the nutrients that the microbes produce.

Soil Science I: 4
The most beneficial, and probably most recognizable macro fauna in soil is the earthworm.
Oligochaeta, a subclass of annelids, or segmented worms, contribute to the texture of soil as we
know it. The earthworms burrow throughout the soil creating tunnels and shafts. These openings
loosen the soil structure providing spaces where other organisms can travel and gain access to
other areas. Water also fills these spaces, adding to the water absorption of the soil, increasing
moisture, which many organisms need, including earthworms. When water gets absorbed into the
ground there is less runoff on the surface. While the worms burrow, they intake soil and dead
plant material. They digest these nutrient-rich materials and produce waste called castings. The
castings contain all of the nutrients that were present in the organic material: nitrogen, potassium,
calcium, phosphorus, and magnesium, and contain microorganisms, such as bacteria and fungi.
The protozoa and nematodes feed on the microorganisms within the castings and release the
nutrients within them. The worm castings are considered to be fertilizers because of their positive
effect on the soil: they help to balance the pH levels, retain moisture, improve drainage, and
control pathogens. Plant nurseries harvest and sell worm castings for this very reason.

Other macro fauna includes nematodes, mollusks, and arthropods. These include a wide
variety of animals that help to maintain a balance in the food web. Some of these animals
consume live or dead plant material, and some are predators of the microorganisms mentioned
above. The larger bugs gain and release some of the nutrients from bacteria when they feed on
them. In addition to this, macro fauna help maintain natural population sizes of their prey. One
very interesting relationship is between macro fauna and bacteria. As the bacteria are eaten, the
more their population grows. This is negative feedback. This is clearly supported in the
relationship between the arthropods and fungi in the following situation. Leaf cutter ants harvest
basidiomycete fungi. They grow the fungi, feeding it leaves, providing it with a desirable living
condition and protection from other organisms. In turn, the ants eat the fungi. The obvious
question begs to be asked, wouldn't the fungi die off if it is constantly being eaten? In actuality,
when it is eaten, it is more stimulated to grow, and it does just that. As long as it is fed (the leaves
provided from the ants) it will continue to grow, and be satisfied living in this situation. This is
the same relationship with bacteria and their predators.

Soil organisms are not always beneficial. organisms that pose a threat to plants and
microorganisms that aid the plants are viewed as pests. Fungi and bacteria have been known for
posing many problems to soil organisms. They can form parasitic relationships with plants,
attacking young or damaged roots and inhibiting their growth. The enzymes that enable these
microorganisms to digest tough plant materials are the same enzymes, such as cellulase, that
allow the bacteria and fungi to penetrate a plant's roots in a pathogenic relationship. Macro fauna
are not faultless. Nematodes can also be parasites, feeding on plants and animals, including
humans. They spread diseases and viruses amongst soil organisms, posing many problems for
farmers and gardeners. Larger nematodes are often introduced as predators to smaller nematodes
to control their population. An arthropod called symphyla, or wire worms, are pests in
greenhouses and gardens. They also feed on plants and microorganisms. Mites pose similar
concerns. As is described above, the organisms in the soil contribute greatly to the wellness of the
plants that grow from the soil. They are dependent on each other and each has a niche to fill to
ensure that the environment stays well-balanced. Organisms do not always behave in a helpful
manner, but pesty organisms occur naturally in an ecosystem. Each type of organism in the soil
plays a role in keeping the soil an ecosystem that is diverse and worthy of our attention.

Soil Science I: 5
LESSON II
BACTERIA: CHARACTERISTICS AND CLASSIFICATION

Bacteria are single celled microbes. The cell structure is simpler than that of other
organisms as there is no nucleus or membrane bound organelles. Instead, their control center
containing the genetic information is contained in a single loop of DNA. Some bacteria have an
extra circle of genetic material called a plasmid. The plasmid often contains genes that give the
bacterium some advantage over other bacteria. For example, it may contain a gene that makes the
bacterium resistant to a certain antibiotic.

Bacteria are tiny, one-celled organisms – generally 4/100,000 of an inch wide (1 µm) and
somewhat longer in length. What bacteria lack in size, they make up in numbers. A teaspoon of
productive soil generally contains between 100 million and 1 billion bacteria. That is as much
mass as two cows per acre.

Bacteria are classified into five groups according to their basic shapes: spherical (cocci),
rod (bacilli), spiral (spirilla), comma (vibrios) or corkscrew (spirochaetes). They can exist as
single cells, in pairs, chains or clusters.

©
ttsz / iStock Different bacterial shapes.

Bacteria are found in every habitat on Earth: soil, rock, oceans and even arctic snow.
Some live in or on other organisms including plants and animals including humans. There are
approximately 10 times as many bacterial cells as human cells in the human body. A lot of these
bacterial cells are found lining the digestive system. Some bacteria live in the soil or on dead plant
matter where they play an important role in the cycling of nutrients. Some types cause food
spoilage and crop damage but others are incredibly useful in the production of fermented foods
such as yoghurt and soy sauce. Relatively few bacteria are parasites or pathogens that cause
disease in animals and plants.

Soil Science I: 6
Bacteria fall into four functional groups. Most are decomposers that consume simple
carbon compounds, such as root exudates and fresh plant litter. By this process, bacteria convert
energy in soil organic matter into forms useful to the rest of the organisms in the soil food web. A
number of decomposers can break down pesticides and pollutants in soil. Decomposers are
especially important in immobilizing, or retaining, nutrients in their cells, thus preventing the loss
of nutrients, such as nitrogen, from the rooting zone.

A second group of bacteria are the mutualists that form partnerships with plants. The most
well-known of these are the nitrogen-fixing bacteria. The third group of bacteria is the pathogens.
Bacterial pathogens include Xymomonas and Erwinia species, and species of Agrobacterium that
cause gall formation in plants. A fourth group, called lithotrophs or chemoautotrophs, obtains its
energy from compounds of nitrogen, sulfur, iron or hydrogen instead of from carbon compounds.
Some of these species are important to nitrogen cycling and degradation of pollutants.

Sample treatment/test of bacteria in the soil and its effect to plant


Courtesy of : https://ars.els-cdn.com/content/image/1-s2.0-S0160412019305410-ga1_lrg.jpg

How do bacteria reproduce?

Bacteria reproduce by binary fission. In this process the bacterium, which is a single cell,
divides into two identical daughter cells. Binary fission begins when the DNA of the bacterium
divides into two (replicates). The bacterial cell then elongates and splits into two daughter cells
each with identical DNA to the parent cell. Each daughter cell is a clone of the parent cell.

Soil Science I: 7
When conditions are favourable such as the right temperature and nutrients are available,
some bacteria like Escherichia coli can divide every 20 minutes. This means that in just seven
hours one bacterium can generate 2,097,152 bacteria. After one more hour the number of bacteria
will have risen to a colossal 16,777,216. That’s why we can quickly become ill when pathogenic
microbes invade our bodies.

Survival Mechanism

Some bacteria can form endospores. These are dormant structures, which are extremely
resistant to hostile physical and chemical conditions such as heat, UV radiation and disinfectants.
This makes destroying them very difficult. Many endospore-producing bacteria are nasty
pathogens, for example Bacillus anthracis, the cause of anthrax.

What Do Bacteria Do?

Bacteria from all four groups perform important services related to water dynamics,
nutrient cycling, and disease suppression. Some bacteria affect water movement by producing
substances that help bind soil particles into small aggregates (those with diameters of 1/10,000-
1/100 of an inch or 2-200µm). Stable aggregates improve water infiltration and the soil’s water-
holding ability. In a diverse bacterial community, many organisms will compete with disease-
causing organisms in roots and on aboveground surfaces of plants.

A Few Important Bacteria

Nitrogen-fixing bacteria form symbiotic associations with the roots of legumes like
clover and lupine, and trees such as alder and locust. Visible nodules are created where bacteria
infect a growing root hair. The plant supplies simple carbon compounds to the bacteria, and the
bacteria convert nitrogen (N2) from air into a form the plant host can use. When leaves or roots
from the host plant decompose, soil nitrogen increases in the surrounding area.

Nitrifying bacteria change ammonium (NH4+) to nitrite (NO2-) then to nitrate (NO3-) –
a preferred form of nitrogen for grasses and most row crops. Nitrate is leached more easily from
the soil, so some farmers use nitrification inhibitors to reduce the activity of one type of nitrifying
bacteria. Nitrifying bacteria are suppressed in forest soils, so that most of the nitrogen remains as
ammonium.

Denitrifying bacteria convert nitrate to nitrogen (N2) or nitrous oxide (N2O) gas.
Denitrifiers are anaerobic, meaning they are active where oxygen is absent, such as in saturated
soils or inside soil aggregates.

Actinomycetes are a large group of bacteria that grow as hyphae like fungi. They are
responsible for the characteristically “earthy” smell of freshly turned, healthy soil. Actinomycetes
decompose a wide array of substrates, but are especially important in degrading recalcitrant (hard-
to-decompose) compounds, such as chitin and cellulose, and are active at high pH levels. Fungi
are more important in degrading these compounds at low pH. A number of antibiotics are
produced by actinomycetes such as Streptomyces.

Soil Science I: 8
Nodules formed where Rhizobium bacteria infected
soybean roots.

Credit: Stephen Temple, New Mexico State University.


Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited)
images.

Rhizobium bacteria nodules

Actinomycetes, such as this Streptomyces, give soil its


"earthy" smell.

Credit: No. 14 from Soil Microbiology and Biochemistry


Slide Set. 1976. J.P. Martin, et al., eds. SSSA, Madison, WI.
Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited)
Actinomycetes
images.

Where Are Bacteria?

Various species of bacteria thrive on different food sources and in different


microenvironments. In general, bacteria are more competitive when labile (easy-to-metabolize)
substrates are present. This includes fresh, young plant residue and the compounds found near
living roots. Bacteria are especially concentrated in the rhizosphere, the narrow region next to and
in the root. There is evidence that plants produce certain types of root exudates to encourage the
growth of protective bacteria.

Bacteria alter the soil environment to the extent that the soil environment will favor certain
plant communities over others. Before plants can become established on fresh sediments, the
bacterial community must establish first, starting with photosynthetic bacteria. These fix
atmospheric nitrogen and carbon, produce organic matter, and immobilize enough nitrogen and
other nutrients to initiate nitrogen cycling processes in the young soil. Then, early successional
plant species can grow. As the plant community is established, different types of organic matter
enter the soil and change the type of food available to bacteria. In turn, the altered bacterial
community changes soil structure and the environment for plants. Some researchers think it may
be possible to control the plant species in a place by managing the soil bacteria community.

Soil Science I: 9
LESSON III
DIFFERENCE BETWEEN ACTINOMYCETES AND FUNGI

Key Difference – Actinomycetes vs Fungi

Microorganisms are tiny organisms which cannot be seen by our naked eyes. There are
several groups of microorganisms. Bacteria and fungi are significant among them. Most bacteria
and fungi are beneficial while a small percentage causes diseases and other harmful effects. Fungi
play various crucial roles in the environment. They the dominant decomposers of organic wastes
and are involved in recycling of nutrients in all terrestrial habitats. Fungi are able to break down
complex material such as cellulose and lignin and help other organisms to absorb nutrients.
Actinomycetes are a group of bacteria which are gram positive and behave like fungi. They are
beneficial in agriculture and soil systems. Actinomycetes grow as colonies which resemble
mycelia of fungi. The key difference between actinomycetes and fungi is that Actinomycetes are
prokaryotic organisms while fungi are eukaryotic organisms.

What are Actinomycetes?


Actinomycetes are a phylum of gram positive bacteria. They are prokaryotic organisms
with a primitive unicellular organization. Actinomycetes are anaerobic microorganisms. They
show filamentous and branching growth pattern on solid substrates resembling fungi mycelia.
Their colonies are extensive like mycelium. Aerial hyphae are found in many genera of
actinomycetes. Some actinomycetes genera are motile and have flagella. Actinomycetes are
responsible for the musty odor ( the smell of freshly ploughed soils) which comes after rain.

Figure 01: Actinomycetes

Soil Science I: 10
Actinomycetes are found in terrestrial and aquatic environments. The common genera of
actinomycetes are Streptomyces, Nocardia, and Micromonospora. Many actinomycetes species
can be observed in the soil. Soil bacteria are harmless to animals and plants. They act as good
decomposers. Hence, they are important in increasing the availability of nutrients for plants.
Actinomycetes produce a wide variety of useful secondary metabolites which have potent
biological activities, including commercially important antibiotics and immunosuppressive
compounds. Some of them are used for manufacturing of commodity chemical, health products,
and agrochemicals.

What are the Similarities Between Actinomycetes and Fungi?

• Actinomycetes and fungi are filamentous.


• Both produce spores.
• Both types are good decomposers.
• Both groups include antibiotic producing species.

Soil Science I: 11
LESSON IV
FUNGI: CHARACTERISTICS AND UNIQUE ADAPTATION
TO SOIL CONDITIONS

Fungi can be single celled or very complex multicellular organisms. They are found in just
about any habitat but most live on the land, mainly in soil or on plant material rather than in sea or
fresh water. A group called the decomposers grow in the soil or on dead plant matter where they
play an important role in the cycling of carbon and other elements. Some are parasites of plants
causing diseases such as mildews, rusts, scabs or canker. In crops fungal diseases can lead to
significant monetary loss for the farmer. A very small number of fungi cause diseases in animals.
In humans these include skin diseases such as athletes’ foot, ringworm and thrush.

Types of fungi

Fungi are subdivided on the basis of their life cycles, the presence or structure of their
fruiting body and the arrangement of and type of spores (reproductive or distributional cells) they
produce.

The three major groups of fungi are:

• Multicellular filamentous moulds.


• Macroscopic filamentous fungi that form large fruiting bodies. Sometimes the group is
referred to as ‘mushrooms’, but the mushroom is just the part of the fungus we see above
ground which is also known as the fruiting body.
• Single celled microscopic yeasts.

© Dennis Kunkel Microscopy / Science Photo Library Candida albicans yeast and hyphae,
coloured scanning electron micrograph (SEM). Great yeast, bud scars and pseudohyphae. A yeast-
like fungus commonly occurring on human skin, in the upper respiratory, alimentary and female
genital tracts. This fungus has a dimorphic life cycle with yeast and hyphal stages. The yeast
produces hyphae (strands) and pseudohyphae. The pseudohyphae can give rise to yeast cells by
apical or lateral budding. Causes candidiasis which includes thrush (an infection of the mouth and
vagina) and vulvo-vaginitis.

Multicellular filamentous molds

Molds are made up of very fine threads (hyphae). Hyphae grow at the tip and divide
repeatedly along their length creating long and branching chains. The hyphae keep growing and
intertwining until they form a network of threads called a mycelium. Digestive enzymes are
secreted from the hyphal tip. These enzymes break down the organic matter found in the soil into
smaller molecules which are used by the fungus as food.

Soil Science I: 12
Some of the hyphal branches grow into the air and spores form on these aerial branches.
Spores are specialised structures with a protective coat that shields them from harsh
environmental conditions such as drying out and high temperatures. They are so small that
between 500 – 1000 could fit on a pin head.

Spores are similar to seeds as they enable the fungus to reproduce. Wind, rain or insects
spread spores. They eventually land in new habitats and if conditions are right, they start to grow
and produce new hyphae. As fungi can’t move, they use spores to find a new environment where
there are fewer competing organisms.

Macroscopic filamentous fungi

Macroscopic filamentous fungi also grow by producing a mycelium below ground. They
differ from moulds because they produce visible fruiting bodies (commonly known as mushrooms
or toadstools) that hold the spores. The fruiting body is made up of tightly packed hyphae which
divide to produce the different parts of the fungal structure, for example the cap and the stem.
Gills underneath the cap are covered with spores and a 10 cm diameter cap can produce up to 100
million spores per hour.

Yeasts

Yeasts are small, lemon-shaped single cells that are about the same size as red blood cells.
They multiply by budding a daughter cell off from the original parent cell. Scars can be seen on
the surface of the yeast cell where buds have broken off. Yeasts such as Saccharomyces play an
important role in the production of bread and in brewing. Yeasts are also one of the most widely
used model organisms for genetic studies, for example in cancer research. Other species of yeast
such as Candida are opportunistic pathogens and cause infections in individuals who do not have
a healthy immune system.

THE LIVING SOIL: FUNGI

Fungi are microscopic cells that usually grow as long threads or strands called hyphae,
which push their way between soil particles, roots, and rocks. Hyphae are usually only several
thousandths of an inch (a few micrometers) in diameter. A single hyphae can span in length from
a few cells to many yards. A few fungi, such as yeast, are single cells.

Hyphae sometimes group into masses called mycelium or thick, cord-like “rhizomorphs” that look
like roots. Fungal fruiting structures (mushrooms) are made of hyphal strands, spores, and some
special structures like gills on which spores form. A single individual fungus can include many
fruiting bodies scattered across an area as large as a baseball diamond.

Fungi perform important services related to water dynamics, nutrient cycling, and disease
suppression. Along with bacteria, fungi are important as decomposers in the soil food web. They
convert hard-to-digest organic material into forms that other organisms can use. Fungal hyphae
physically bind soil particles together, creating stable aggregates that help increase water
infiltration and soil water holding capacity.

Soil Science I: 13
Soil fungi can be grouped into three general functional groups based on how they get their energy.

➢ Decomposers – saprophytic fungi – convert dead organic material into fungal biomass, carbon
dioxide (CO2), and small molecules, such as organic acids. These fungi generally use complex
substrates, such as the cellulose and lignin, in wood, and are essential in decomposing the
carbon ring structures in some pollutants. A few fungi are called “sugar fungi” because they
use the same simple substrates as do many bacteria. Like bacteria, fungi are important for
immobilizing, or retaining, nutrients in the soil. In addition, many of the secondary metabolites
of fungi are organic acids, so they help increase the accumulation of humic-acid rich organic
matter that is resistant to degradation and may stay in the soil for hundreds of years.

➢ Mutualists – the mycorrhizal fungi – colonize plant roots. In exchange for carbon from the
plant, mycorrhizal fungi help solubolize phosphorus and bring soil nutrients (phosphorus,
nitrogen, micronutrients, and perhaps water) to the plant. One major group of mycorrhizae, the
ectomycorrhizae (see third photo below), grow on the surface layers of the roots and are
commonly associated with trees. The second major group of mycorrhizae are the
endomycorrhizae that grow within the root cells and are commonly associated with grasses,
row crops, vegetables, and shrubs. Arbuscular mycorrhizal (AM) fungi are a type of
endomycorrhizal fungi. Ericoid mycorrhizal fungi can by either ecto- or endomycorrhizal.

➢ The third group of fungi, pathogens or parasites, cause reduced production or death when
they colonize roots and other organisms. Root-pathogenic fungi, such as Verticillium, Pythium,
and Rhizoctonia, cause major economic losses in agriculture each year. Many fungi help
control diseases. For example, nematode-trapping fungi that parasitize disease-causing
nematodes, and fungi that feed on insects may be useful as biocontrol agents.

Many plants depend on fungi to help extract nutrients from


the soil. Tree roots (brown) are connected to the symbiotic
mycorrhizal structure (bright white) and fungal hyphae (thin
white strands) radiating into the soil.

Credit: Randy Molina, Oregon State University, Corvallis. Please contact the
Soil and Water Conservation Society at pubs@swcs.org for assistance with
Mycorrhizal Structure and Fungal Hyphae copyrighted (credited) images.

Fungus beginning to decompose leaf veins in grass clippings.

Credit: No. 48 from Soil Microbiology and Biochemistry Slide Set. 1976. J.P.
Martin, et al., eds. SSSA, Madison WI. Please contact the Soil and Water
Conservation Society at pubs@swcs.org for assistance with copyrighted
(credited) images.

Fungus Decomposing Leaf Veins

Soil Science I: 14
Ectomycorrhizae are important for nutrient absorption by
tree and grape roots. The fungus does not actually invade
root cells but forms a sheath that penetrates between plant
cells. The sheath in this photo is white, but they may be
black, orange, pink, or yellow.

Credit: USDA, Forest Service, PNW Research Station, Corvallis, Oregon. Please
contact the Soil and Water Conservation Society at pubs@swcs.org for assistance
with copyrighted (credited) images.
Ectomycorrhizae Fungi

The dark, round masses inside the cells of this clover root are
vesicules for the arbuscular mycorrhizal fungus (AM).

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation
Society at pubs@swcs.org for assistance with copyrighted (credited) images.

Vesicular-arbuscular Mycorrhizal Fungus

Where Are Fungi?

Saprophytic fungi are commonly active around woody plant residue. Fungal hyphae have
advantages over bacteria in some soil environments. Under dry conditions, fungi can bridge gaps
between pockets of moisture and continue to survive and grow, even when soil moisture is too
low for most bacteria to be active. Fungi are able to use nitrogen up from the soil, allowing them
to decompose surface residue which is often low in nitrogen.

Fungi are aerobic organisms. Soil which becomes anaerobic for significant periods
generally loses its fungal component. Anaerobic conditions often occur in waterlogged soil and in
compacted soils.

Fungi are especially extensive in forested lands. Forests have been observed to increase in
productivity as fungal biomass increases.

In arid rangeland systems, such as southwestern deserts, fungi


pipe scarce water and nutrients to plants.

Credit: Jerry Barrow, USDA-ARS Jornada Experimental Range, Las Cruces, NM.
Please contact the Soil and Water Conservation Society at pubs@swcs.org for
assistance with copyrighted (credited) images.

Arid Rangeland System

Soil Science I: 15
Mushrooms, common in forest systems, are the fruiting bodies
made by a group of fungi called basidiomycetes. Mushrooms are
"the tip of the iceberg" of an extensive network of underground
hyphae.

Credit: Ann Lewandowski, NRCS Soil Quality Institute. Please contact the Soil and
Water Conservation Society at pubs@swcs.org for assistance with copyrighted
(credited) images.
Basidiomycete Fungi Mushroom

Mycorrhizal Fungi in Agriculture

Mycorrhiza is a symbiotic association between fungi and plant roots and is unlike either
fungi or roots alone. Most trees and agricultural crops depend on or benefit substantially from
mycorrhizae. The exceptions are many members of the Cruciferae family (e.g., broccoli,
mustard), and the Chenopodiaceae family (e.g. lambsquarters, spinach, beets), which do not form
mycorrhizal associations. The level of dependency on mycorrhizae varies greatly among varieties
of some crops, including wheat and corn.

Land management practices affect the formation of mycorrhizae. The number of


mycorrhizal fungi in soil will decline in fallowed fields or in those planted to crops that do not
form mycorrhizae. Frequent tillage may reduce mycorrhizal associations, and broad-spectrum
fungicides are toxic to mycorrhizal fungi. Very high levels of nitrogen or phosphorus fertilizer
may reduce inoculation of roots. Some inoculums of mycorrhizal fungi are commercially
available and can be added to the soil at planting time.

Mycorrhizal fungi link root cells to soil particles; sand


grains are bound to a root by hyphae from endophytes
(fungi similar to mycorrhizae), and by polysaccharides
secreted by the plant and the fungi.

Credit: Jerry Barrow, USDA-ARS Jornada Experimental Range, Las


Cruces, NM. Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images.

Mycorrhizal Fungi Link Root to Soil Particles

Soil Science I: 16
Soil Science I: 17
LESSON V
OTHER ORGANISMS: PROTOZOA, NEMATODES, ARTHROPODS AND WORMS,

1. PROTOZOA

Protozoa are single celled organisms. They come in many different shapes and sizes ranging
from an Amoeba which can change its shape to Paramecium with its fixed shape and complex
structure. They live in a wide variety of moist habitats including fresh water, marine environments
and the soil.

Some are parasitic, which means they live in other plants and animals including humans, where
they cause disease. Plasmodium, for example, causes malaria. They are motile and can move by:

➢ Cilia - tiny hair like structures that cover the outside of the microbe. They beat in a regular
continuous pattern like flexible oars.
➢ Flagella - long thread-like structures that extend from the cell surface. The flagella move in
a whip-like motion that produces waves that propel the microbe around.
➢ Amoeboid movement - the organism moves by sending out pseudopodia, temporary
protrusions that fill with cytoplasm that flows from the body of the cell.

The Living Soil: Protozoa

Protozoa are single-celled animals that feed primarily on bacteria, but also eat other
protozoa, soluble organic matter, and sometimes fungi. They are several times larger than bacteria
- ranging from 1/5000 to 1/50 of an inch (5 to 500 µm) in diameter. As they eat bacteria, protozoa
release excess nitrogen that can then be used by plants and other members of the food web.

Protozoa are classified into three groups based on their shape: Ciliates are the largest and
move by means of hair-like cilia. They eat the other two types of protozoa, as well as bacteria.
Amoebae also can be quite large and move by means of a temporary foot or "pseudopod."
Amoebae are further divided into testate amoebae (which make a shell-like covering) and naked
amoebae (without a covering). Flagellates are the smallest of the protozoa and use a few whip-
like flagella to move.

Protozoa play an important role in nutrient cycling by feeding


intensively on bacteria. Notice the size of the speck-like
bacteria next to the oval protozoa and large, angular sand
particle.

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images.

Protozoa Feed on Bacteria

Soil Science I: 18
Bacteria ingested by an amoeba.

Credit: No. 35 from Soil Microbiology and Biochemistry Slide Set. 1976. J.P.
Martin, et al., eds. SSSA, Madison, WI. Please contact the Soil and Water
Conservation Society at pubs@swcs.org for assistance with copyrighted (credited)
images.

Bacteria Ingested by an Amoeba

Flagellates have one or two flagella which they use to propel or


pull their way through soil. A flagellum can be seen extending
from the protozoan on the left. The tiny specks are bacteria.

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images.

Flagellate Protozoa

Ciliates are the largest of the protozoa and the least numerous.
They consume up to ten thousand bacteria per day, and release
plant available nitrogen. Ciliates use the fine cilia along their
bodies like oars to move rapidly through soil.

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images.

Ciliates

What Do Protozoa Do?

Protozoa play an important role in mineralizing nutrients, making them available for use
by plants and other soil organisms. Protozoa (and nematodes) have a lower concentration of
nitrogen in their cells than the bacteria they eat. (The ratio of carbon to nitrogen for protozoa is
10:1 or much more and 3:1 to 10:1 for bacteria.) Bacteria eaten by protozoa contain too much
nitrogen for the amount of carbon protozoa need. They release the excess nitrogen in the form of
ammonium (NH4+). This usually occurs near the root system of a plant. Bacteria and other
organisms rapidly take up most of the ammonium, but some is used by the plant. (See figure
below for explanation of mineralization and immobilzation.)

Another role that protozoa play is in regulating bacteria populations. When they graze on
bacteria, protozoa stimulate growth of the bacterial population (and, in turn, decomposition rates
and soil aggregation.) Exactly why this happens is under some debate, but grazing can be thought

Soil Science I: 19
of like pruning a tree - a small amount enhances growth, too much reduces growth or will modify
the mix of species in the bacterial community.

Protozoa are also an important food source for other soil organisms and help to suppress disease
by competing with or feeding on pathogens.

Where Are Protozoa?

Protozoa need bacteria to eat and water in which to move, so moisture plays a big role in
determining which types of protozoa will be present and active. Like bacteria, protozoa are
particularly active in the rhizosphere next to roots.

Typical numbers of protozoa in soil vary widely - from a thousand per teaspoon in low
fertility soils to a million per teaspoon in some highly fertile soils. Fungal-dominated soils (e.g.
forests) tend to have more testate amoebae and ciliates than other types. In bacterial-dominated
soils, flagellates and naked amoebae predominate. In general, high clay-content soils contain a
higher number of smaller protozoa (flagellates and naked amoebae), while coarser textured soils
contain more large flagellates, amoebae of both varieties, and ciliates.

Nematodes and Protozoa

Protozoa and bacterial-feeding nematodes compete for their common food resource:
bacteria. Some soils have high numbers of either nematodes or protozoa, but not both. The
significance of this difference to plants is not known. Both groups consume bacteria and release
NH4+.

Bug Biography: Soil Dwelling Vampires

Fungi-eating Protozoa

Most protozoa eat bacteria, but one group of amoebae, the vampyrellids, eat fungi. The
perfectly round holes drilled through the fungal cell wall, much like the purported puncture marks
on the neck of a vampire's victim, are evidence of the presence of vampyrellid amoebae. The
amoebae attach to the surface of fungal hyphae and generate enzymes that eat through the fungal
cell wall. The amoeba then sucks dry or engulfs the cytoplasm inside the fungal cell before
moving on to its next victim.

Vampyrellids attack many fungi including root pathogens, such as Gaeumannomyces


graminis, shown in the photo. This fungus attacks wheat roots and causes take-all disease.

Soil Science I: 20
2. NEMATODES

Nematode, also called roundworm, any worm of the phylum Nematoda. Nematodes are
among the most abundant animals on Earth. They occur as parasites in animals and plants or as
free-living forms in soil, fresh water, marine environments, and even such unusual places as
vinegar, beer malts, and water-filled cracks deep within Earth’s crust. The number of named
species is about 20,000, but it is probable that only a small proportion of the free-living forms
have been identified. A great deal of research has been conducted on the parasitic forms because
most of them have some medical, veterinary, or economic importance.

Nematodes are bilaterally symmetrical, elongate, and usually tapered at both ends. Some
species possess a pseudocoel, a fluid-filled body cavity between the digestive tract and the body
wall. Like arthropods and members of six other phyla, nematodes secrete an external cuticle that
is periodically molted. These animals have been provisionally grouped together as the Ecdysozoa,
a taxonomic category based on the assumption that molting has evolved only once. So far, gene
sequence data from several molecules support such an assumption.

The Living Soil: Nematodes

Nematodes are non-segmented worms typically 1/500 of an inch (50 µm) in diameter and
1/20 of an inch (1 mm) in length. Those few species responsible for plant diseases have received a
lot of attention, but far less is known about the majority of the nematode community that plays
beneficial roles in soil.

An incredible variety of nematodes function at several trophic levels of the soil food web. Some
feed on the plants and algae (first trophic level); others are grazers that feed on bacteria and fungi
(second trophic level); and some feed on other nematodes (higher trophic levels).

Free-living nematodes can be divided into four broad groups based on their diet.

• Bacterial-feeders consume bacteria.


• Fungal-feeders feed by puncturing the cell wall of fungi and sucking out the internal
contents.
• Predatory nematodes eat all types of nematodes and protozoa. They eat smaller
organisms whole, or attach themselves to the cuticle of larger nematodes, scraping away
until the prey’s internal body parts can be extracted.
• Omnivores eat a variety of organisms or may have a different diet at each life stage. Root-
feeders are plant parasites, and thus are not free-living in the soil.

Most nematodes in the soil are not plant parasites. Beneficial


nematodes help control disease and cycle nutrients.

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation Society
at pubs@swcs.org for assistance with copyrighted (credited) images.

Beneficial Nematodes

Soil Science I: 21
A predatory nematode consumes a smaller nematode.

Credit: Kathy Merrifield, Oregon State University, Corvallis. Please contact the
Soil and Water Conservation Society at pubs@swcs.org for assistance with
copyrighted (credited) images.

Predatory Nematode

What Do Nematodes Do?

Nutrient cycling. Like protozoa, nematodes are important in mineralizing, or releasing, nutrients
in plant-available forms. When nematodes eat bacteria or fungi, ammonium (NH4+) is released
because bacteria and fungi contain much more nitrogen than the nematodes require.

Grazing. At low nematode densities, feeding by nematodes stimulates the growth rate of prey
populations. That is, bacterial-feeders stimulate bacterial growth, plant-feeders stimulate plant
growth, and so on. At higher densities, nematodes will reduce the population of their prey. This
may decrease plant productivity, may negatively impact mycorrhizal fungi, and can reduce
decomposition and immobilization rates by bacteria and fungi. Predatory nematodes may regulate
populations of bacterial-and fungal-feeding nematodes, thus preventing over-grazing by those
groups. Nematode grazing may control the balance between bacteria and fungi, and the species
composition of the microbial community.

Dispersal of microbes. Nematodes help distribute bacteria and fungi through the soil and along
roots by carrying live and dormant microbes on their surfaces and in their digestive systems.

Food source. Nematodes are food for higher level predators, including predatory nematodes, soil
microarthropods, and soil insects. They are also parasitized by bacteria and fungi.

Disease suppression and development. Some nematodes cause disease. Others consume disease-
causing organisms, such as root-feeding nematodes, or prevent their access to roots. These may be
potential biocontrol agents.

Fungal-feeding nematodes have small, narrow stylets, or spears,


in their stoma (mouth) which they use to puncture thecell walls
of fungal hyphae and withdraw the cell fluid. This interaction
releases plant-available nitrogen from fungal biomass.

Credit: Elaine R. Ingham. Please contact the Soil and Water


Conservation Society at pubs@swcs.org for assistance with
copyrighted (credited) images.
Fungal-feeding Nematode

Soil Science I: 22
This bacterial-feeding nematode, Elaphonema, has ornate lip
structures that distinguish it from other nematodes. Bacterial-
feeders release plant-available nitrogen when they consume
bacteria.

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images .
Bacterial-feeding Nematode

The Pratylenchus, or lesion nematode, has a shorter, thicker


stylet in its mouth than the root feeder below.

Credit: Kathy Merrifield, Oregon State University, Corvallis. Please contact the Soil
and Water Conservation Society at pubs@swcs.org for assistance with copyrighted
(credited) images.

Lesion Nematode

Root-feeding nematodes use their stylets to puncture the thick


cell wall of plant root cells and siphon off the internal contents
of the plant cell. This usually causes economically significant
damage to crops. The curved stylet seen inside this nematode is
characteristic of the genus Trichodorus.

Credit: Elaine R. Ingham. Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images.
Root-feeding Nematode

Where Are Nematodes?

Nematodes are concentrated near their prey groups. Bacterial-feeders abound near roots
where bacteria congregate; fungal-feeders are near fungal biomass; root-feeders are concentrated
around roots of stressed or susceptible plants. Predatory nematodes are more likely to be abundant
in soils with high numbers of nematodes.

Because of their size, nematodes tend to be more common in coarser-textured soils.


Nematodes move in water films in large (>1/500 inch or 50 µm) pore spaces.

Agricultural soils generally support less than 100 nematodes in each teaspoon (dry gram)
of soil. Grasslands may contain 50 to 500 nematodes, and forest soils generally hold several
hundred per teaspoon. The proportion of bacterial-feeding and fungal-feeding nematodes is
related to the amount of bacteria and fungi in the soil. Commonly, less disturbed soils contain
more predatory nematodes, suggesting that predatory nematodes are highly sensitive to a wide
range of disturbances.

Soil Science I: 23
Nematodes and Water Quality

Nematodes may be useful indicators of soil quality because of their tremendous diversity
and their participation in many functions at different levels of the soil food web. Several
researchers have proposed approaches to assessing the status of soil quality by counting the
number of nematodes in different families or trophic groups.* In addition to their diversity,
nematodes may be useful indicators because their populations are relatively stable in response to
changes in moisture and temperature (in contrast to bacteria), yet nematode populations respond
to land management changes in predictable ways. Because they are quite small and live in water
films, changes in nematode populations reflect changes in soil microenvironments.

*Blair, J. M. et al. 1996. Soil invertebrates as indicators of soil quality. In Methods for Assessing Soil Quality, SSSA Special
Publication 49, pp. 273-291.

Bug Biography: Nematode Trappers

One group of fungi may be a useful biological control agent against parasitic nematodes. These
predatory fungi grow through the soil, setting out traps when they detect signs of their prey. Some
species use sticky traps, others make circular rings of hyphae to constrict their prey. When the
trap is set, the fungi put out a lure, attracting nematodes that are looking for lunch. The nematode,
however, becomes lunch for the fungus.

3. ARTHROPODS

Most arthropods are small animals. Only aquatic forms are able to attain substantial sizes,
because their bodies are supported in part by the surrounding water. The extinct chelicerate
Eurypterida, for example, reached a length of 1.8 metres (5.9 feet), and some modern spider crabs
may weigh up to 6.4 kilograms (14 pounds) and span 3.8 metres or more. Terrestrial arthropods
do not grow very large. The largest adult insects and spiders do not weigh more than 100 grams
(0.22 pound); however, there is evidence that larvae of Megasoma actaeon, a type of rhinoceros
beetle, can sometimes exceed 200 grams (0.44 pound). The beetle Goliathus regius measures 15
centimetres (5.9 inches) in length and 10 centimetres in width, while the butterfly Ornithoptera
victoriae of the Solomon Islands has a wing span exceeding 30 centimetres (about 1 foot). One of
the longest insects is the phasmid (walkingstick) Phryganistria chinensis, a specimen of which
measured 62.4 centimetres (about 2 feet) in length. The phasmid Phobaeticus chani reaches a
length of more than 30 centimetres. The smallest arthropods include some parasitic wasps, beetles
of the family Ptiliidae, and mites that are less than 0.25 millimetre (0.01 inch) in length, despite
their complex structures.

The Living Soil: Arthropods

Many bugs, known as arthropods, make their home in the soil. They get their name from their
jointed (arthros) legs (podos). Arthropods are invertebrates, that is, they have no backbone, and
rely instead on an external covering called an exoskeleton.

Soil Science I: 24
The 200 species of mites in this microscope view were
extracted from one square foot of the top two inches of
forest litter and soil. Mites are poorly studied, but
enormously significant for nutrient release in the soil.

Credit: Val Behan-Pelletier, Agriculture and Agri-Food Canada. Please


contact the Soil and Water Conservation Society at pubs@swcs.org for
assistance with copyrighted (credited) images.

Microscopic View of Mites

Arthropods range in size from microscopic to several inches in length. They include
insects, such as springtails, beetles, and ants; crustaceans such as sowbugs; arachnids such as
spiders and mites; myriapods, such as centipedes and millipedes; and scorpions.

Nearly every soil is home to many different arthropod species. Certain row-crop soils
contain several dozen species of arthropods in a square mile. Several thousand different species
may live in a square mile of forest soil.

Arthropods can be grouped as shredders, predators, herbivores, and fungal-feeders, based


on their functions in soil. Most soil-dwelling arthropods eat fungi, worms, or other arthropods.
Root-feeders and dead-plant shredders are less abundant. As they feed, arthropods aerate and mix
the soil, regulate the population size of other soil organisms, and shred organic material.

➢ Shredders

Many large arthropods frequently seen on the soil surface are shredders. Shredders chew
up dead plant matter as they eat bacteria and fungi on the surface of the plant matter. The most
abundant shredders are millipedes and sowbugs, as well as termites, certain mites, and roaches. In
agricultural soils, shredders can become pests by feeding on live roots if sufficient dead plant
material is not present.

Millipedes are also called Diplopods because they possess


two pairs of legs on each body segment. They are generally
harmless to people, but most millipedes protect themselves
from predators by spraying an offensive odor from their
skunk glands. This desert-dwelling giant millipede is about
8 inches long.
Millipedes are Shredders
Orthoporus ornatus.

Credit: David B. Richman, New Mexico State University, Las Cruces. Please
contact the Soil and Water Conservation Society at pubs@swcs.org for
assistance with copyrighted (credited) images.

Soil Science I: 25
Sowbugs are relatives of crabs and lobsters. Their powerful
mouth-parts are used to fragment plant residue and leaf
litter.

Credit: Gerhard Eisenbeis and Wilfried Wichard. 1987. Atlas on the Biology of
Soil Arthropods. Springer-Verlag, New York. P. 111. Please contact the Soil
and Water Conservation Society at pubs@swcs.org for assistance with
copyrighted (credited) images.

Sowbug

➢ Predators

Predators and micropredators can be either generalists, feeding on many different prey
types, or specialists, hunting only a single prey type. Predators include centipedes, spiders,
ground-beetles, scorpions, skunk-spiders, pseudoscorpions, ants, and some mites. Many predators
eat crop pests, and some, such as beetles and parasitic wasps, have been developed for use as
commercial biocontrols.

This 1/8 of an inch long spider lives near the soil surface where
it attacks other soil arthropods. The spider's eyes are on the tip
of the projection above its head.
Walckenaera acuminata.

Credit: Gerhard Eisenbeis and Wilfried Wichard. 1987. Atlas on the Biology of Soil
Arthropods. Springer-Verlag, New York. P. 23. Please contact the Soil and Water
Spider - Walckenaera acuminata Conservation Society at pubs@swcs.org for assistance with copyrighted (credited)
images.

The wolf-spider wanders around as a solitary hunter. The


mother wolf-spider carries her young to water and feeds them
by regurgitation until they are ready to hunt on their own.

Credit: Trygve Steen, Portland State University, Portland, Oregon. Please contact
the Soil and Water Conservation Society at pubs@swcs.org for assistance with
copyrighted (credited) images.
Wolf-spider

The pseudoscorpion looks like a baby scorpion, except it has no


tail. It produces venom from glandsin its claws and silk from its
mouth parts. It lives in the soil and leaf litter of grasslands,
forests, deserts and croplands. Some hitchhike under the wings
of beetles.

Credit: David B. Richman, New Mexico State University, Las Cruces. Please contact
the Soil and Water Conservation Society at pubs@swcs.org for assistance with
Pseudoscorpion
copyrighted (credited) images.

Soil Science I: 26
Long, slim centipedes crawl through spaces in the soil preying
on earthworms and other soft-skinned animals. Centipede
species with longer legs are familiar around homes and in leaf
litter.

Credit: No. 40 from Soil Microbiology and Biochemistry Slide Set. 1976. J.P.
Martin, et al., eds. SSSA, Madison, WI. Please contact the Soil and Water
Conservation Society at pubs@swcs.org for assistance with copyrighted (credited)
images.
Centipede

Predatory mites prey on nematodes, springtails, other mites,


and the larvae of insects. This mite is 1/25 of an inch (1mm)
long. Pergamasus sp.

Credit: Gerhard Eisenbeis and Wilfried Wichard. 1987. Atlas on the Biology of Soil
Arthropods. Springer-Verlag, New York. P. 83. Please contact the Soil and Water
Conservation Society at pubs@swcs.org for assistance with copyrighted (credited)
images.

Predatory Mite

The powerful mouthparts on the tiger beetle (a carabid beetle)


make it a swift and deadly ground-surface predator. Many
species of carabid beetles are common in cropland.

Credit: Cicindela campestris. D.I. McEwan/Aguila Wildlife Images. Please contact


the Soil and Water Conservation Society at pubs@swcs.org for assistance with
Tiger Beetle copyrighted (credited) images.

Rugose harvester ants are scavengers rather than predators.


They eat dead insects and gather seeds in grasslands and
deserts where they burrow 10 feet into the ground. Their sting
is 100 times more powerful than a fire ant sting.
Pogonomyrmex rugosus.

Credit: David B. Richman, New Mexico State University, Las Cruces. Please contact
the Soil and Water Conservation Society at pubs@swcs.org for assistance with
Rugose Harvester Ants copyrighted (credited) images.

Soil Science I: 27
➢ Herbivores

Numerous root-feeding insects, such as cicadas, mole-crickets, and anthomyiid flies (root-
maggots), live part of all of their life in the soil. Some herbivores, including rootworms and
symphylans, can be crop pests where they occur in large numbers, feeding on roots or other plant
parts.

The symphylan, a relative of the centipede, feeds on plant roots and


can become a major crop pest if its population is not controlled by
other organisms.

Credit: Ken Gray Collection, Department of Entomology, Oregon


State University, Corvallis. Please contact the Soil and Water
Conservation Society at pubs@swcs.org for assistance with
copyrighted (credited) images.
Symphylan Feeds on Plant Roots

➢ Fungal Feeders

Arthropods that graze on fungi (and to some extent bacteria) include most springtails,
some mites, and silverfish. They scrape and consume bacteria and fungi off root surfaces. A large
fraction of the nutrients available to plants is a result of microbial-grazing and nutrient release by
fauna.

This pale-colored and blind springtail is typical of fungal-


feeding springtails that live deep in the surface layer of natural
and agricultural soils throughout the world.

Credit: Andrew R. Moldenke, Oregon State University, Corvallis. Please contact the
Soil and Water Conservation Society at pubs@swcs.org for assistance with
Fungal-feeding Springtail copyrighted (credited) images.

Oribatid turtle-mites are among the most numerous of the


micro-arthropods. This millimeter-long species feeds on fungi.
Euzetes globulus.

Credit: Gerhard Eisenbeis and Wilfried Wichard. 1987. Atlas on the Biology of Soil
Arthropods. Springer-Verlag, New York. P. 103. Please contact the Soil and Water
Conservation Society at pubs@swcs.org for assistance with copyrighted (credited)
images.

Oribatid turtle-mite

What Is In Your Soil?

If you would like to see what kind of organisms are in your soil, you can easily make a
pitfall trap to catch large arthropods, and a Burlese funnel to catch small arthropods.

Soil Science I: 28
Make a pitfall trap by sinking a pint- or quart-sized container (such as a yogurt cup) into
the ground so the rim is level with the soil surface. If desired, fashion a roof over the cup to keep
the rain out, and add 1/2 of an inch of non-hazardous antifreeze to the cup to preserve the
creatures and prevent them from eating one another. Leave in place for a week and wait for soil
organisms to fall into the trap.

To make a Burlese funnel, set a piece of 1/4 inch rigid wire screen in the bottom of a
funnel to support the soil. (A funnel can be made by cutting the bottom off a plastic soda bottle.)
Half fill the funnel with soil, and suspend it over a cup with a bit of anti-freeze or ethyl alcohol in
the bottom as a preservative.

Suspend a light bulb about 4 inches over the soil to drive the organisms out of the soil and
into the cup. Leave the light bulb on for about 3 days to dry out the soil. Then pour the alcohol
into a shallow dish and use a magnifying glass to examine the organisms.

What Do Arthropods Do?

Although the plant feeders can become pests, most arthropods perform beneficial
functions in the soil-plant system.

Shred organic material. Arthropods increase the surface area accessible to microbial attack by
shredding dead plant residue and burrowing into coarse woody debris. Without shredders, a
bacterium in leaf litter would be like a person in a pantry without a can-opener – eating would be
a very slow process. The shredders act like can-openers and greatly increase the rate of
decomposition. Arthropods ingest decaying plant material to eat the bacteria and fungi on the
surface of the organic material.

Stimulate microbial activity. As arthropods graze on bacteria and fungi, they stimulate the
growth of mycorrhizae and other fungi, and the decomposition of organic matter. If grazer
populations get too dense the opposite effect can occur – populations of bacteria and fungi will
decline. Predatory arthropods are important to keep grazer populations under control and to
prevent them from over-grazing microbes.

Mix microbes with their food. From a bacterium’s point-of-view, just a fraction of a millimeter
is infinitely far away. Bacteria have limited mobility in soil and a competitor is likely to be closer
to a nutrient treasure. Arthropods help out by distributing nutrients through the soil, and by
carrying bacteria on their exoskeleton and through their digestive system. By more thoroughly
mixing microbes with their food, arthropods enhance organic matter decomposition.

Mineralize plant nutrients. As they graze, arthropods mineralize some of the nutrients in
bacteria and fungi, and excrete nutrients in plant-available forms.

Enhance soil aggregation. In most forested and grassland soils, every particle in the upper
several inches of soil has been through the gut of numerous soil fauna. Each time soil passes
through another arthropod or earthworm, it is thoroughly mixed with organic matter and mucus
and deposited as fecal pellets. Fecal pellets are a highly concentrated nutrient resource, and are a
mixture of the organic and inorganic substances required for growth of bacteria and fungi. In
many soils, aggregates between 1/10,000 and 1/10 of an inch (0.0025mm and 2.5mm) are actually
fecal pellets.

Soil Science I: 29
Burrow. Relatively few arthropod species burrow through the soil. Yet, within any soil
community, burrowing arthropods and earthworms exert an enormous influence on the
composition of the total fauna by shaping habitat. Burrowing changes the physical properties of
soil, including porosity, water-infiltration rate, and bulk density.

Stimulate the succession of species. A dizzying array of natural bio-organic chemicals permeates
the soil. Complete digestion of these chemicals requires a series of many types of bacteria, fungi,
and other organisms with different enzymes. At any time, only a small subset of species is
metabolically active – only those capable of using the resources currently available. Soil
arthropods consume the dominant organisms and permit other species to move in and take their
place, thus facilitating the progressive breakdown of soil organic matter.

Control pests. Some arthropods can be damaging to crop yields, but many others that are present
in all soils eat or compete with various root- and foliage-feeders. Some (the specialists) feed on
only a single type of prey species. Other arthropods (the generalists), such as many species of
centipedes, spiders, ground-beetles, rove-beetles, and gamasid mites, feed on a broad range of
prey. Where a healthy population of generalist predators is present, they will be available to deal
with a variety of pest outbreaks. A population of predators can only be maintained between pest
outbreaks if there is a constant source of non-pest prey to eat. That is, there must be a healthy and
diverse food web.

A fundamental dilemma in pest control is that tillage and insecticide application have enormous
effects on non- target species in the food web. Intense land use (especially monoculture, tillage,
and pesticides) depletes soil diversity. As total soil diversity declines, predator populations drop
sharply and the possibility for subsequent pest outbreaks increases.

Where Do Arthropods Live?

The abundance and diversity of soil fauna diminishes significantly with soil depth. The
great majority of all soil species are confined to the top three inches. Most of these creatures have
limited mobility, and are probably capable of “cryptobiosis,” a state of “suspended animation”
that helps them survive extremes of temperature, wetness, or dryness that would otherwise be
lethal.

As a general rule, larger species are active on the soil surface, seeking temporary refuge
under vegetation, plant residue, wood, or rocks. Many of these arthropods commute daily to
forage within herbaceous vegetation above, or even high in the canopy of trees. (For instance, one
of these tree-climbers is the caterpillar-searcher used by foresters to control gypsy moth). Some
large species capable of true burrowing live within the deeper layers of the soil.

Below about two inches in the soil, fauna are generally small – 1/250 to 1/10 of an inch.
(Twenty-five of the smallest of these would fit in a period on this page.) These species are usually
blind and lack prominent coloration. They are capable of squeezing through minute pore spaces
and along root channels. Sub-surface soil dwellers are associated primarily with the rhizosphere
(the soil volume immediately adjacent to roots).

Abundance of Arthropods

A single square yard of soil will contain 500 to 200,000 individual arthropods, depending
upon the soil type, plant community, and management system. Despite these large numbers, the
biomass of arthropods in soil is far less than that of protozoa and nematodes.

Soil Science I: 30
In most environments, the most abundant soil dwellers are springtails and mites, though
ants and termites predominate in certain situations, especially in desert and tropical soils. The
largest number of arthropods are in natural plant communities with few earthworms (such as
conifer forests). Natural communities with numerous earthworms (such as grassland soils) have
the fewest arthropods. Apparently, earthworms out-compete arthropods, perhaps by excessively
reworking their habitat or eating them incidentally. However, within pastures and farm lands
arthropod numbers and diversity are generally thought to increase as earthworm populations rise.
Burrowing earthworms probably create habitat space for arthropods in agricultural soils.

Bug Biography: Springtails

Springtail
Springtails are the most abundant arthropods in many agricultural and rangeland soils. populations
of tens of thousands per square yard are frequent. When foraging, springtails walk with 3 pairs of
legs like most insects, and hold their tail tightly tucked under the belly. If attacked by a predator,
body fluid rushes into the tail base, forcing the tail to slam down and catapult the springtail as
much as a yard away. Springtails have been shown to be beneficial to crop plants by releasing
nutrients and by feeding upon diseases caused by fungi.

4. WORMS

Worm, any of various unrelated invertebrate animals that typically have soft, slender,
elongated bodies. Worms usually lack appendages; polychaete annelids are a conspicuous
exception. Worms are members of several invertebrate phyla, including Platyhelminthes
(flatworms), Annelida (segmented worms), Nemertea (ribbon worms), Nematoda (roundworms,
pinworms, etc.), Sipuncula (peanutworms), Echiura (spoonworms), Acanthocephala (spiny-
headed worms), Pogonophora (beardworms), and Chaetognatha (arrowworms).

The term is also loosely applied to centipedes and millipedes; to larval (immature) forms
of other invertebrates, particularly those of certain insects; and to some vertebrates—e.g., the
blindworm (Anguis fragilis), a limbless, snakelike lizard. At one time all phyla of wormlike
animals were classed as Vermes, a term no longer in common use.

The major groups of worms include various species of flatworm, annelid, ribbon worm,
spiny-headed worm, and aschelminth (qq.v.). Worms typically have an elongated, tubelike body,
usually rather cylindrical, flattened, or leaflike in shape and often without appendages. They vary
in size from less than 1 mm (0.04 inch) in certain nematodes to more than 30 m (100 feet) in
certain ribbon worms (phylum Nemertea).Worms are universal in distribution, occurring in
marine, freshwater, and terrestrial habitats. Some types of worms are parasitic, others are free-
living. From a human perspective, worms are important as soil conditioners (e.g., annelids,

Soil Science I: 31
aschelminths) and as parasites of people and domestic animals (e.g., platyhelminths,
aschelminths) and of crops (e.g., aschelminths). Ecologically, worms form an important link in
the food chains in virtually all ecosystems of the world.

The Living Soil: Earthworms

Of all the members of the soil food web, earthworms need the least introduction. Most people
become familiar with these soft, slimy, invertebrates at a young age. Earthworms are
hermaphrodites, meaning that they exhibit both male and female characteristics.

They are major decomposers of dead and decomposing organic matter, and derive their nutrition
from the bacteria and fungi that grow upon these materials. They fragment organic matter and
make major contributions to recycling the nutrients it contains.

Earthworms occur in most temperate soils and many tropical soils. They are divided into 23
families, more than 700 genera, and more than 7,000 species. They range from an inch to two
yards in length and are found seasonally at all depths in the soil.

In terms of biomass and overall activity, earthworms dominate the world of soil invertebrates,
including arthropods.

Earthworms generate tons of casts per acre each year, dramatically


altering soil structure.

Credit: Clive A. Edwards, The Ohio State University, Columbus.


Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited) images.
Earthworms

A corn leaf pulled into a night crawler burrow.

Credit: Soil and Water Management Research Unit, USDA-


Agricultural Research Service, St. Paul, Minnesota. Please contact the
Soil and Water Conservation Society at pubs@swcs.org for assistance
with copyrighted (credited) images.

Corn Leaf Pulled into Night


Crawler Burrow

What Do Earthworms Do?

Earthworms dramatically alter soil structure, water movement, nutrient dynamics, and plant
growth. They are not essential to all healthy soil systems, but their presence is usually an indicator
of a healthy system. Earthworms perform several beneficial functions.

Stimulate microbial activity. Although earthworms derive their nutrition from microorganisms,
many more microorganisms are present in their feces or casts than in the organic matter that they
consume. As organic matter passes through their intestines, it is fragmented and inoculated with
microorganisms. Increased microbial activity facilitates the cycling of nutrients from organic
matter and their conversion into forms readily taken up by plants.

Soil Science I: 32
Mix and aggregate soil. As they consume organic matter and mineral particles, earthworms
excrete wastes in the form of casts, a type of soil aggregate. Charles Darwin calculated that
earthworms can move large amounts of soil from the lower strata to the surface and also carry
organic matter down into deeper soil layers. A large proportion of soil passes through the guts of
earthworms, and they can turn over the top six inches (15 cm) of soil in ten to twenty years.

Increase infiltration. Earthworms enhance porosity as they move through the soil. Some species
make permanent burrows deep into the soil. These burrows can persist long after the inhabitant
has died, and can be a major conduit for soil drainage, particularly under heavy rainfall. At the
same time, the burrows minimize surface water erosion. The horizontal burrowing of other
species in the top several inches of soil increases overall porosity and drainage.

Improve water-holding capacity. By fragmenting organic matter, and increasing soil porosity
and aggregation, earthworms can significantly increase the water-holding capacity of soils.

Provide channels for root growth. The channels made by deep-burrowing earthworms are lined
with readily available nutrients and make it easier for roots to penetrate deep into the soil.

Bury and shred plant residue. Plant and crop residue are gradually buried by cast material
deposited on the surface and as earthworms pull surface residue into their burrows.

A mixture of soil and organic matter within an earthworm


burrow. Earthworms incorporate large amounts of organic
matter into the soil.

Credit: Clive A. Edwards, The Ohio State University,


Columbus. Please contact the Soil and Water
Earthworm Burrow with Organic Matter
Conservation Society at pubs@swcs.org for assistance
with copyrighted (credited) images.

Some worms live in permanent vertical burrows such as


these. Others move horizontally near the surface, filling
their burrow with casts as they move.

Credit: North Appalachian Experimental Watershed,


USDA-Agricultural Research Service, Coshocton, Ohio.
Vertical Earthworm Burrow
Please contact the Soil and Water Conservation Society
at pubs@swcs.org for assistance with copyrighted
(credited) images.

Earthworm Burrow graphic

Soil Science I: 33
Where Are Earthworms?

Different species of earthworms inhabit different parts of the soil and have distinct feeding
strategies. They can be separated into three major ecological groups based on their feeding and
burrowing habits. All three groups are common and important to soil structure.

Surface soil and litter species – Epigeic species. These species live in or near surface plant litter.
They are typically small and are adapted to the highly variable moisture and temperature
conditions at the soil surface. The worms found in compost piles are epigeic and are unlikely to
survive in the low organic matter environment of soil.

Upper soil species – Endogeic species. Some species move and live in the upper soil strata and
feed primarily on soil and associated organic matter (geophages). They do not have permanent
burrows, and their temporary channels become filled with cast material as they move through the
soil, progressively passing it through their intestines.

Deep-burrowing species – Anecic species. These earthworms, which are typified by the “night
crawler,” Lumbricus terrestris, inhabit more or less permanent burrow systems that may extend
several meters into the soil. They feed mainly on surface litter that they pull into their burrows.
They may leave plugs, organic matter, or cast (excreted soil and mineral particles) blocking the
mouth of their burrows.

Looking for Earthworms?

It is easy to determine whether you have an adequate population of earthworms in your


soil. Look for their casts in the forms of little piles of soil, mineral particles, or organic matter at
the soil surface. They can be seen moving over the soil surface or even breeding, particularly on
warm, damp nights. Dump a spade full of moist soil into a bucket or onto a sheet of plastic, and
sort through for earthworms. Can you identify different species? To find the deep burrowing
species, pour a dilute mustard solution onto the soil. Many will quickly come to the soil surface in
response to this irritant.

Abundance and Distribution of Earthworms

The majority of temperate and many tropical soils support significant earthworm
populations. A square yard of cropland in the United States can contain from 50-300 earthworms,
or even larger populations in highly organic soils. A similar area of grassland or temperate
woodlands will have from 100-500 earthworms. Based on their total biomass, earthworms are the
predominant group of soil invertebrates in most soils.

The family of earthworms that is most important in enhancing agricultural soil is


Lumbricidae, which includes the genuses Lumbricus, Aporrectodea, and several others.
Lumbricids originated in Europe and have been transported by human activities to many parts of
the world. The United States has only one or two known native species of lumbricids. Others were
brought to this country by settlers (probably in potted plants from Europe), and were distributed
down the waterways.

Generally, lumbricids are much more common in the north and east than in the drier south
and west of the United States. They tend to be more abundant in loam and clay loam and even in
silty soil, than in sandy soil and heavy clay. Populations also build up in irrigated soil. Earthworm

Soil Science I: 34
populations tend to increase with soil organic matter levels and decrease with soil disturbances,
such as tillage and potentially harmful chemicals.

Casts at the soil surface are evidence that earthworms are


shredding, mixing, and burying surface residue.

Credit: Soil and Water Management Research Unit, USDA-


Agricultural Research Service, St. Paul, Minnesota. Please
contact the Soil and Water Conservation Society at
Earthworm Casts pubs@swcs.org for assistance with copyrighted (credited)
images.

This earthworm burrow is an opening in an otherwise crusted soil


surface.

Credit: Clive A. Edwards, The Ohio State University, Columbus.


Please contact the Soil and Water Conservation Society at
pubs@swcs.org for assistance with copyrighted (credited)
Earthworm Burrow Crusted Soil Surface images.

Interactions of Earthworms with Other Members of the Food Web

The lives of earthworms and microbes are closely intertwined. Earthworms derive their
nutrition from fungi, bacteria, and possibly protozoa and nematodes, and they promote the activity
of these organisms by shredding and increasing the surface area of organic matter and making it
more available to small organisms.

Earthworms also influence other soil-inhabiting invertebrates by changing the amount and
distribution of organic matter and microbial populations. There is good evidence that earthworm
activity affects the spatial distribution of soil microarthropod communities in the soil.

Earthworms have few invertebrate enemies, other than flatworms and a species of parasitic
fly. Their main predators are a wide range of birds and mammals that prey upon them at the soil
surface.

Earthworms and Water Quality

Earthworms improve water infiltration and water holding capacity because their
shredding, mixing, and defecating enhances soil structure. In addition, burrows provide quick
entry for water into and through soil. High infiltration rates help prevent pollution by minimizing
runoff, erosion, and chemical transport to surface waters.

There is concern that burrows may increase the transport of pollutants, such as nitrates or
pesticides, into groundwater. However, the movement of potential pollutants through soil is not a
straightforward process and it is not clear when earthworm activity will or will not have a
negative impact on groundwater quality.

Soil Science I: 35
Whether pollutants reach groundwater depends on a number of factors, including the
location of pollutants on the surface or within soil, the quantity and intensity of rain, how well
water moves into and through other parts of the soil, and characteristics of the burrows. The
horizontal burrows of endogeic earthworms (such as Aporrectodea tuberculata, which are
common in Midwestern fields) do not transport water and solutes as deeply as the vertical
burrows of night crawlers (L. terrestris) and other anecic species. Even vertical burrows, however,
are not direct channels for water movement. They have bends and turns and are lined with organic
matter that adsorbs many potential pollutants from the water.

Although there is much more to learn about how earthworms affect water movement
through soil, they clearly help minimize pollution of surface waters by improving infiltration rates
and decreasing runoff.

A mound of organic matter was moved aside to expose the


entrance to a burrow. L. terrestris will quickly replug its burrow if
its mound is removed.

Credit: North Appalachian Experimental Watershed, USDA-


Agricultural Research Service, Coshocton, Ohio. Please contact
Earthworm Burrow Entrance
the Soil and Water Conservation Society at pubs@swcs.org for
assistance with copyrighted (credited) images.

L. terrestris mating, and earthworm cocoons. Earthworms mate


periodically throughout the year, except when environmental
conditions are unfavorable. The worms form slime tubes to help
adhere to each other during copulation which may take as long as
an hour.
Earthworms Mating

After the worms separate, they each produce a cocoon. One or


two worms will hatch from a cocoon after several weeks. L.
terrestris cocoons are about a quarter inch long.

Credit: Clive A. Edwards, The Ohio State University, Columbus.


Please contact the Soil and Water Conservation Society at
Earthworm Cocoons pubs@swcs.org for assistance with copyrighted (credited)
images.

Bug Biography: Night Crawlers and Tillage

The substitution of conventional tillage by no-till or conservation tillage is increasingly


common and widely adopted in the United States and elsewhere. In these situations, earthworms,
particularly the “night crawler,” Lumbricus terrestris L., are especially important. Earthworms
become the main agent for incorporating crop residue into the soil by pulling some into their
burrows and by slowly burying the remainder under casts laid on the soil surface.

Soil Science I: 36
In reduced tillage systems, surface residue builds up and triggers growth in earthworm
populations. Earthworms need the food and habitat provided by surface residue, and they eat the
fungi that become more common in no-till soils. As earthworm populations increase, they pull
more and more residue into their burrows, helping to mix organic matter into the soil, improving
soil structure and water infiltration.

LESSON VI
BENEFICIAL ACTIVITIES OF SOIL ORGANISMS

Soil organism, any organism inhabiting the soil during part or all of its life. Soil
organisms, which range in size from microscopic cells that digest decaying organic material to
small mammals that live primarily on other soil organisms, play an important role in maintaining
fertility, structure, drainage, and aeration of soil. They also break down plant and animal tissues,
releasing stored nutrients and converting them into forms usable by plants. Some soil organisms
are pests. Among the soil organisms that are pests of crops are nematodes, slugs and snails,
symphylids, beetle larvae, fly larvae, caterpillars, and root aphids. Some soil organisms cause rots,
some release substances that inhibit plant growth, and others are hosts for organisms that cause
animal diseases.Since most of the functions of soil organisms are beneficial, earth with large
numbers of organisms in it tends to be fertile; one square metre of rich soil can harbour as many
as 1,000,000,000 organisms.Soil organisms are commonly divided into five arbitrary groups
according to size, the smallest of which are the protists—including bacteria, actinomycetes, and
algae. Next are the microfauna, which are less than 100 microns in length and generally feed upon
other microorganisms. The microfauna include single-celled protozoans, some smaller flatworms,
nematodes, rotifers, and tardigrades (eight-legged invertebrates). The mesofauna are somewhat
larger and are heterogeneous, including creatures that feed on microorganisms, decaying matter,
and living plants. The category includes nematodes, mites, springtails (wingless insects so called
for the springing organ which enables them to leap), the insectlike proturans, which feed on fungi,
and the pauropods.

The fourth group, the macrofauna, are also quite diverse. The most common example is
the potworm, a white, segmented worm that feeds on fungi, bacteria, and decaying plant material.
The group also includes slugs, snails, and millipedes, which feed on plants, and centipedes,
beetles and their larvae, and the larvae of flies, which feed on other organisms or on decaying
matter.

Megafauna constitute the largest soil organisms and include the largest earthworms,
perhaps the most important creatures that live in the topsoil. Earthworms pass both soil and
organic matter through their guts, in the process aerating the soil, breaking up the litter of organic
material on its surface, and moving material vertically from the surface to the subsoil. This is
extremely important to soil fertility, and it develops the structure of the soil as a matrix for plants
and other organisms. It has been estimated that earthworms completely turn over the equivalent of
all the soil on the planet to a depth one inch (2.5 cm) every 10 years. Some vertebrates are also in
the megafauna category; these include all sorts of burrowing animals, such as snakes, lizards,
gophers, badgers, rabbits, hares, mice, and moles.

One of the most important roles of soil organisms is breaking up the complex substances
in decaying plants and animals so that they can be used again by living plants. This involves soil
organisms as catalysts in a number of natural cycles, among the most prominent being the carbon,
nitrogen, and sulfur cycles.

Soil Science I: 37
The carbon cycle begins in plants, which combine carbon dioxide from the atmosphere
with water to make plant tissues such as leaves, stems, and fruits. Animals eat the plants and
convert the tissues into animal tissues. The cycle is completed when the animals die and their
decaying tissues are eaten by soil organisms, a process that releases carbon dioxide.

Proteins are the basic stuff of organic tissues, and nitrogen is an essential element of all
proteins. The availability of nitrogen in forms that plants can use is a basic determinant of the
fertility of soils; the role of soil organisms in facilitating the nitrogen cycle is therefore of great
importance. When a plant or animal dies, soil organisms break up the complex proteins,
polypeptides, and nucleic acids in their bodies and produce ammonium, ions, nitrates, and nitrites
that plants then use to build their body tissues.

BOth bacteria and blue-green algae can fix nitrogen directly from the atmosphere, but this
is less vital to plant development than the symbiotic relationship between the bacteria genus
Rhizobium and leguminous plants and certain trees and shrubs. In return for secretions from their
host that encourage their growth and multiplication, Rhizobia fix nitrogen in nodules of the host
plant’s roots, providing nitrogen in a form usable by the plant.

Soil organisms also participate in the sulfur cycle, mostly by breaking up the naturally
abundant sulfur compounds in the soil so that this vital element is available to plants. The smell of
rotten eggs so common in swamps and marshes is due to the hydrogen sulfide produced by these
microorganisms.

Though soil organisms have become less important in agriculture due to the development
of synthetic fertilizers, they play a vital role in woodlands, especially in the creation of humus, a
finely separated complex of organic materials composed of decaying leaves and other vegetable
matter.

When a leaf falls it cannot be eaten by most animals. After the water-soluble components
of the leaf are leached out, fungi and other microflora attack its structure, making it soft and
pliable. Now the litter is palatable to a wide variety of invertebrates, which fragment it into a
mulch. The multipedes, wood lice, fly larvae, springtails, and earthworms leave the litter
relatively unchanged organically, but they create a suitable substrate for the growth of the primary
decomposers that break it into simpler chemical compounds. There is also a group called
secondary decomposers (some creatures, such as the springtails, are in both groups), which break
it down even further.

So the organic matter of leaves is constantly being digested and redigested by waves of
increasingly smaller organisms. Eventually the humic substance that remains may be as little as
one-fourth of the original organic matter of the litter. Gradually this humus is mixed into the soil
by burrowing animals (such as moles, rabbits, etc.) and by the action of the earthworms.

Though some soil organisms can become pests—especially when a single crop is grown
repeatedly in the same field, encouraging the proliferation of organisms that prey on their roots—
by and large they are essential elements in the process of life, death, and decay, which rejuvenates
the environment.

Soil Science I: 38
LESSON VII
ROLE OF SOIL ORGANIC MATTER

Once a land manager begins working towards enhancing soil organic matter, a series of soil
changes and environmental benefits follow. The rate and degree of these changes and the best
suite of practices needed to achieve results vary with soil and climate. Initially, managing for
greater soil organic matter may require higher pesticide, herbicide, or nutrient applications. In
time, productivity and environmental quality will be enhanced.

1. Apply practices that enhance soil organic matter

• Diverse, high biomass crop rotations


• Cover crops
• Reduced tillage
• Rotational or prescribed grazing

2. Organic matter dynamics change

• Increased surface residue forms a physical barrier to wind and water erosion.
• Higher residue rotations and cover crops contribute more organic matter and nutrients to
the soil.
• Less soil disturbance means lower organic matter losses.

3. Soil properties change

• Surface structure becomes more stable and less prone to crusting and erosion.
• Water infiltration increases and runoff decreases when soil structure improves.
• Soil organic matter holds 10 to 1,000 times more water and nutrients than the same
amount of soil minerals.
• Beneficial soil organisms become more numerous and active with diverse crop rotations
and higher organic matter levels.

4. Air quality, water quality, and agricultural productivity improve

• Dust, allergens, and pathogens in the air immediately decline.


• Sediment and nutrient loads decline in surface water as soon as soil aggregation increases
and runoff decreases.
• Ground and surface water quality improve because better structure, infiltration, and
biological activity make soil a more effective filter.
• Crops are better able to withstand drought when infiltration and water holding capacity
increase.
• Organic matter may bind pesticides, making them less active. Soils managed for organic
matter may suppress disease organisms, which could reduce pesticide needs.
• Crop health and vigor increase when soil biological activity and diversity increase.
• Wildlife habitat improves when residue management improves.

Soil Science I: 39
LESSON VIII

ORGANIC MATTER DECOMPOSITION AND THE SOIL FOOD WEB

Soil Organic Matter

When plant residues are returned to the soil, various organic compounds undergo
decomposition. Decomposition is a biological process that includes the physical breakdown and
biochemical transformation of complex organic molecules of dead material into simpler organic
and inorganic molecules (Juma, 1998).

The continual addition of decaying plant residues to the soil surface contributes to the
biological activity and the carbon cycling process in the soil. Breakdown of soil organic matter
and root growth and decay also contribute to these processes. Carbon cycling is the continuous
transformation of organic and inorganic carbon compounds by plants and micro- and macro-
organisms between the soil, plants and the atmosphere (Figure 2)

Soil Science I: 40
FIGURE 2
Carbon cycle

Decomposition of organic matter is largely a biological process that occurs naturally. Its
speed is determined by three major factors: soil organisms, the physical environment and the
quality of the organic matter (Brussaard, 1994). In the decomposition process, different products
are released: carbon dioxide (CO2), energy, water, plant nutrients and resynthesized organic
carbon compounds. Successive decomposition of dead material and modified organic matter
results in the formation of a more complex organic matter called humus (Juma, 1998). This
process is called humification. Humus affects soil properties. As it slowly decomposes, it colours
the soil darker; increases soil aggregation and aggregate stability; increases the CEC (the ability to
attract and retain nutrients); and contributes N, P and other nutrients.

Soil organisms, including micro-organisms, use soil organic matter as food. As they break
down the organic matter, any excess nutrients (N, P and S) are released into the soil in forms that
plants can use. This release process is called mineralization. The waste products produced by
micro-organisms are also soil organic matter. This waste material is less decomposable than the
original plant and animal material, but it can be used by a large number of organisms. By breaking
down carbon structures and rebuilding new ones or storing the C into their own biomass, soil
biota plays the most important role in nutrient cycling processes and, thus, in the ability of a soil
to provide the crop with sufficient nutrients to harvest a healthy product. The organic matter
content, especially the more stable humus, increases the capacity to store water and store
(sequester) C from the atmosphere.

The soil food web

The soil ecosystem (Box 1) can be defined as an interdependent life-support system


composed of air, water, minerals, organic matter, and macro- and micro-organisms, all of which
function together and interact closely.

The organisms and their interactions enhance many soil ecosystem functions and make up
the soil food web. The energy needed for all food webs is generated by primary producers: the
plants, lichens, moss, photosynthetic bacteria and algae that use sunlight to transform CO2 from

Soil Science I: 41
the atmosphere into carbohydrates. Most other organisms depend on the primary producers for
their energy and nutrients; they are called consumers.

BOX 1
Some functions of a healthy soil ecosystem

• Decompose organic matter towards humus.


• Retain N and other nutrients.
• Glue soil particles together for best structure.
• Protect roots from diseases and parasites.
• Make retained nutrients available to the plant.
• Produce hormones that help plants grow.
• Retain water.

Soil life plays a major role in many natural processes that determine nutrient and water
availability for agricultural productivity. The primary activities of all living organisms are
growing and reproducing. By-products from growing roots and plant residues feed soil organisms.
In turn, soil organisms support plant health as they decompose organic matter, cycle nutrients,
enhance soil structure and control the populations of soil organisms, both beneficial and harmful
(pests and pathogens) in terms of crop productivity.

The living part of soil organic matter includes a wide variety of micro-organisms such as
bacteria, viruses, fungi, protozoa and algae. It also includes plant roots, insects, earthworms, and
larger animals such as moles, mice and rabbits that spend part of their life in the soil. The living
portion represents about 5 percent of the total soil organic matter. Micro-organisms, earthworms
and insects help break down crop residues and manures by ingesting them and mixing them with
the minerals in the soil, and in the process recycling energy and plant nutrients. Sticky substances
on the skin of earthworms and those produced by fungi and bacteria help bind particles together.
Earthworm casts are also more strongly aggregated (bound together) than the surrounding soil as
a result of the mixing of organic matter and soil mineral material, as well as the intestinal mucus
of the worm. Thus, the living part of the soil is responsible for keeping air and water available,
providing plant nutrients, breaking down pollutants and maintaining the soil structure.

The composition of soil organisms depends on the food source (which in turn is season
dependent). Therefore, the organisms are neither uniformly distributed through the soil nor
uniformly present all year. However, in some cases their biogenic structures remain. Each species
and group exists where it can find appropriate food supply, space, nutrients and moisture (Plate
2). Organisms occur wherever organic matter occurs (Ingham, 2000). Therefore, soil organisms
are concentrated: around roots, in litter, on humus, on the surface of soil aggregates and in spaces
between aggregates. For this reason, they are most prevalent in forested areas and cropping
systems that leave a lot of biomass on the surface.

Soil Science I: 42
Plate 2
Termites create their own living conditions near their
preferred food sources. Inside the colony life is highly organized.

The activity of soil organisms follows seasonal as well as daily patterns. Not all organisms
are active at the same time. Most are barely active or even dormant. Availability of food is an
important factor that influences the level of activity of soil organisms and thus is related to land
use and management (Figure 3). Practices that increase numbers and activity of soil organisms
include: no tillage or minimal tillage; and the maintenance of plant and annual residues that
reduce disturbance of soil organisms and their habitat and provide a food supply.

Different groups of organisms can be distinguished in the soil (Brussaard and Juma,
1995). Table 1 classifies them by size. Table 2 classifies them by function.

Decomposition process

Fresh residues consist of recently deceased micro-organisms, insects and earthworms, old
plant roots, crop residues, and recently added manures.

Crop residues contain mainly complex carbon compounds originating from cell walls
(cellulose, hemicellulose, etc.). Chains of carbon, with each carbon atom linked to other carbons,
form the “backbone” of organic molecules. These carbon chains, with varying amounts of
attached oxygen, H, N, P and S, are the basis for both simple sugars and amino acids and more
complicated molecules of long carbon chains or rings. Depending on their chemical structure,
decomposition is rapid (sugars, starches and proteins), slow (cellulose, fats, waxes and resins) or
very slow (lignin).

Soil Science I: 43
Figure 3
Fluctuations in microbial biomass at different stages of crop development in
conventional agriculture compared with systems with residue retention and high organic
matter input

Source: Balota, 1996

TABLE 1
Classification of soil organisms

Micro-organisms Microflora <5 µm Bacteria


Fungi
Microfauna <100 µm Protozoa
Nematodes
Macro-organisms Meso-organisms 100 µm - 2 mm Springtails
Mites
Macro-organisms 2 - 20 mm Earthworms
Millipedes
Woodlice
Snails and slugs
Plants Algae 10 µm
Roots > 10 µm

Note: Clay particles are smaller than 2 µm.


Source: adapted from Swift, Heal and Anderson, 1979.

Soil Science I: 44
TABLE 2
Essential functions performed by different members of soil organisms (biota)

Functions Organisms involved


Maintenance of soil Bioturbating invertebrates and plant roots, mycorrhizae and
structure some other micro-organisms
Regulation of soil Most bioturbating invertebrates and plant roots
hydrological processes
Gas exchange and carbon Mostly micro-organisms and plant roots, some C protected
sequestration in large compact biogenic invertebrate aggregates
(accumulation in soil)
Soil detoxification Mostly micro-organisms
Nutrient cycling Mostly micro-organisms and plant roots, some soil- and
litter-feeding invertebrates
Decomposition of organic Various saprophytic and litter-feeding invertebrates
matter (detritivores), fungi, bacteria, actinomycetes and other
micro-organisms
Suppression of pests, Plants, mycorrhizae and other fungi, nematodes, bacteria and
parasites and diseases various other micro-organisms, collembola, earthworms,
various predators
Sources of food and Plant roots, various insects (crickets, beetle larvae, ants,
medicines termites), earthworms, vertebrates, micro-organisms and
their by-products
Symbiotic and asymbiotic Rhizobia, mycorrhizae, actinomycetes, diazotrophic bacteria
relationships with plants and various other rhizosphere micro-organisms, ants
and their roots
Plant growth control Direct effects: plant roots, rhizobia, mycorrhizae,
(positive and negative) actinomycetes, pathogens, phytoparasitic nematodes,
rhizophagous insects, plant-growth promoting rhizosphere
micro-organisms, biocontrol agents Indirect effects: most
soil biota

During the decomposition process, microorganisms convert the carbon structures of fresh
residues into transformed carbon products in the soil. There are many different types of organic
molecules in soil. Some are simple molecules that have been synthesized directly from plants or
other living organisms. These relatively simple chemicals, such as sugars, amino acids, and
cellulose are readily consumed by many organisms. For this reason, they do not remain in the soil
for a long time. Other chemicals such as resins and waxes also come directly from plants, but are
more difficult for soil organisms to break down.

Humus is the result of successive steps in the decomposition of organic matter. Because of
the complex structure of humic substances, humus cannot be used by many micro-organisms as an
energy source and remains in the soil for a relatively long time.

Soil Science I: 45
Non-humic substances: significance and function

Non-humic organic molecules are released directly from cells of fresh residues, such as
proteins, amino acids, sugars, and starches. This part of soil organic matter is the active, or easily
decomposed, fraction. This active fraction is influenced strongly by weather conditions, moisture
status of the soil, growth stage of the vegetation, addition of organic residues, and cultural
practices, such as tillage. It is the main food supply for various organisms in the soil.

Carbohydrates occur in the soil in three main forms: free sugars in the soil solution,
cellulose and hemicellulose; complex polysaccharides; and polymeric molecules of various sizes
and shapes that are attached strongly to clay colloids and humic substances (Stevenson, 1994).
The simple sugars, cellulose and hemicellulose, may constitute 5-25 percent of the organic matter
in most soils, but are easily broken down by micro-organisms.

Polysaccharides (repeating units of sugar-type molecules connected in longer chains)


promote better soil structure through their ability to bind inorganic soil particles into stable
aggregates. Research indicates that the heavier polysaccharide molecules may be more important
in promoting aggregate stability and water infiltration than the lighter molecules (Elliot and
Lynch, 1984). Some sugars may stimulate seed germination and root elongation. Other soil
properties affected by polysaccharides include CEC, anion retention and biological activity.

The soil lipids form a very diverse group of materials, of which fats, waxes and resins
make up 2-6 percent of soil organic matter. The significance of lipids arises from the ability of
some compounds to act as growth hormones. Others may have a depressing effect on plant
growth.

Soil N occurs mainly (> 90 percent) in organic forms as amino acids, nucleic acids and
amino sugars. Small amounts exist in the form of amines, vitamins, pesticides and their
degradation products, etc. The rest is present as ammonium (NH 4-) and is held by the clay
minerals.

Compounds and function of humus

Humus or humified organic matter is the remaining part of organic matter that has been
used and transformed by many different soil organisms. It is a relatively stable component formed
by humic substances, including humic acids, fulvic acids, hymatomelanic acids and humins (Tan,
1994). It is probably the most widely distributed organic carbon-containing material in terrestrial
and aquatic environments. Humus cannot be decomposed readily because of its intimate
interactions with soil mineral phases and is chemically too complex to be used by most
organisms. It has many functions (Box 2).

One of the most striking characteristics of humic substances is their ability to interact with
metal ions, oxides, hydroxides, mineral and organic compounds, including toxic pollutants, to
form water-soluble and water-insoluble complexes. Through the formation of these complexes,
humic substances can dissolve, mobilize and transport metals and organics in soils and waters, or
accumulate in certain soil horizons. This influences nutrient availability, especially those nutrients
present at microconcentrations only (Schnitzer, 1986). Accumulation of such complexes can
contribute to a reduction of toxicity, e.g. of aluminium (Al) in acid soils (Tan and Binger, 1986),
or the capture of pollutants - herbicides such as Atrazine or pesticides such as Tefluthrin - in the
cavities of the humic substances (Vermeer, 1996).

Soil Science I: 46
Humic and fulvic substances enhance plant growth directly through physiological and
nutritional effects. Some of these substances function as natural plant hormones (auxines and
gibberillins) and are capable of improving seed germination, root initiation, uptake of plant
nutrients and can serve as sources of N, P and S (Tan, 1994; Schnitzer, 1986). Indirectly, they
may affect plant growth through modifications of physical, chemical and biological properties of
the soil, for example, enhanced soil water holding capacity and CEC, and improved tilth and
aeration through good soil structure (Stevenson, 1994).

About 35-55 percent of the non-living part of organic matter is humus. It is an important
buffer, reducing fluctuations in soil acidity and nutrient availability. Compared with simple
organic molecules, humic substances are very complex and large, with high molecular weights.
The characteristics of the well-decomposed part of the organic matter, the humus, are very
different from those of simple organic molecules. While much is known about their general
chemical composition, the relative significance of the various types of humic materials to plant
growth is yet to be established.

Humus consists of different humic substances:

• Fulvic acids: the fraction of humus that is soluble in water under all pH conditions. Their
colour is commonly light yellow to yellow-brown.
• Humic acids: the fraction of humus that is soluble in water, except for conditions more
acid than pH 2. Common colours are dark brown to black.
• Humin: the fraction of humus that is not soluble in water at any pH and that cannot be
extracted with a strong base, such as sodium hydroxide (NaOH). Commonly black in
colour.

The term acid is used to describe humic materials because humus behaves like weak acids.

Fulvic and humic acids are complex mixtures of large molecules. Humic acids are larger than
fulvic acids. Research suggests that the different substances are differentiated from each other on
the basis of their water solubility.

Fulvic acids are produced in the earlier stages of humus formation. The relative amounts of
humic and fulvic acids in soils vary with soil type and management practices. The humus of forest
soils is characterized by a high content of fulvic acids, while the humus of agricultural and
grassland areas contains more humic acids.

BOX 2
Humic substances retain nutrients available on demand for plants

Functions of humus:

• improved fertilizer efficiency;


• longlife N - for example, urea performs 60-80 days longer;
• improved nutrient uptake, particularly of P and Ca;
• stimulation of beneficial soil life;
• provides magnified nutrition for reduced disease, insect and frost impact;
• salinity management - humates “buffer” plants from excess sodium;
• organic humates are a catalyst for increasing soil C levels.

Soil Science I: 47
LESSON IX

TRANSFORMATION OF NITROGEN

Nitrogen cycle, circulation of nitrogen in various forms through nature. Nitrogen, a


component of proteins and nucleic acids, is essential to life on Earth. Although 78 percent by
volume of the atmosphere is nitrogen gas, this abundant reservoir exists in a form unusable by
most organisms. Through a series of microbial transformations, however, nitrogen is made
available to plants, which in turn ultimately sustain all animal life. The steps, which are not
altogether sequential, fall into the following classifications: nitrogen fixation, nitrogen
assimilation, ammonification, nitrification, and denitrification.

Nitrogen fixation, in which nitrogen gas is converted into inorganic nitrogen compounds,
is mostly (90 percent) accomplished by certain bacteria and blue-green algae. A much smaller
amount of free nitrogen is fixed by abiotic means (e.g., lightning, ultraviolet radiation, electrical
equipment) and by conversion to ammonia through the Haber-Bosch process.

Nitrates and ammonia resulting from nitrogen fixation are assimilated into the specific
tissue compounds of algae and higher plants. Animals then ingest these algae and plants,
converting them into their own body compounds.

The remains of all living things—and their waste products—are decomposed by


microorganisms in the process of ammonification, which yields ammonia (NH3) and ammonium
(NH4+). (Under anaerobic, or oxygen-free, conditions, foul-smelling putrefactive products may
appear, but they too are converted to ammonia in time.) Ammonia can leave the soil or be
converted into other nitrogen compounds, depending in part on soil conditions. Nitrification, a
process carried out by nitrifying bacteria, transforms soil ammonia into nitrates (NO3−), which
plants can incorporate into their own tissues. Nitrates also are metabolized by denitrifying
bacteria, which are especially active in water-logged anaerobic soils. The action of these bacteria
tends to deplete soil nitrates, forming free atmospheric nitrogen.

Soil Science I: 48
Suggested Viewing, Readings and References
Required Videos for viewing:

1. NITROGEN CYCLE | Biology Animation


https://www.youtube.com/watch?v=njzFtvMPJ3A
2. CARBON CYCLE | Biology Animation
https://www.youtube.com/watch?v=fcna9slTljs
3. Soil Biology and Organic Matter - Ray Weil
https://www.youtube.com/watch?v=rAZnbxLnTXU
4. Understanding and Managing Soil Biology for Soil Health and Crop Production
https://www.youtube.com/watch?v=JGQP--xtGVc

Sources and Online References:


1. https://www.sare.org/wp-content/uploads/Conservation-Tillage-Systems-in-the-
Southeast_compressed.pdf
2. https://www.nrcs.usda.gov/wps/portal/nrcs/detailfull/soils/health/biology/?cid=nrcs142p2_
053863
3. https://www.fao.org/3/a0100e/a0100e04.htm
4. Agro/Hort 100 Into to Plants. http://molbio.nmsu.edu/hort100/soils.html [accessed August
2006]
5. Biology 501 Class Notes. August 1, 2006. University of Pennsylvania.
6. http://www.mansfield.ohio-state.edu/~sabedon/biol3035.htm
7. http://mypeoplepc.com/members/arbra/trinity/id14.html
8. http://nsm1.utdallas.edu/bio/Robinson/fall2005/Fungi_Fall05.htm
9. http://www.soilhealth.com/index.htm
10. Ingham, E. (2000). Soil Biology Primer [online]. Available:
soils.usda.gov/sqi/concepts/soil_biology/index.html [accessed August, 2006].
11. Jackson, R. and Raw, F. (1966). Life in the Soil. New York, St. Martin's Press.
12. Krogh, D. (2005). Biology: A Guide to the Natural World. 3rd ed. New Jersey, Pearon
Prentice Hall.
13. Mackenzie, F. (2003). Our Changing Planet: An Introduction to earth System Science and
Global Environmental Change. 3rd ed. Prentice Hall.
14. National Science and Technology Center: Soil Biological Communities.
http://www.blm.gov/nstc/soil/index.html
15. https://www.differencebetween.com/difference-between-actinomycetes-and-vs-fungi/
16. www.wikipedia.org

Soil Science I: 49
Activity 5
Synthesis Paper

Instruction: Do the readings of content of the given link below.


Make a synthesis paper base from the readings
(see instructions for the format of assignment and activity).
1. https://www.fao.org/3/a0100e/a0100e06.htm#TopOfPage
2. https://www.fao.org/3/a0100e/a0100e07.htm#TopOfPage
3. https://www.fao.org/3/a0100e/a0100e08.htm#TopOfPage

Reaction Paper 5

Soil organism and Organic Matter


Instruction: Do access this links below and make some random readings and make at least 500
words essay of your major takeaways (knowledge acquired/learnings)

https://reliefweb.int/sites/reliefweb.int/files/resources/Soil_Report_Main_001.pdf
(note: in doing reaction paper use only the front page of assignment/activity format and remove
the rubric)

Soil Science I: 50

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