Who lives in the soil ?? The Furrow Soil is an archive of life on earth! Quantifying Biodiversity Group Known Species Estimated Total Species Percentage Known Vascular plants 220,000 270,000 81% Algae 40,000 60,000 67% Fungi 69,000 1,500,000 5% 3,000 30,000 10% Bacteria Most occur in soil (Hawksworth, 1991) Most are tiny !!! Soil organisms vary widely in size Body size increasing Abundance Viruses Bacteria Fungi Microflora Algae Protozoa Microfauna Nematodes Microarthropods Mesofauna Enchytraeids Earthworms Macrofauna Ants, termites, spiders Mollusks Megafauna Others: rodents, snakes, voles, amphibians, etc. Body sizes and shapes are adaptations to the many different spatial niches in soil BACTERIA cocci have many different shapes bacillus spirilla filamentou s SSS A Typically > 10 billion bacteria per gram of soil What do bacteria do in soil ? Colonize aerobic and anaerobic environments Decompose labile substrates Mediate redox transformations Nourish bacterivores Fix N N-fixing nodules on a cowpea plant Legume love affair Sarrantonio FUNGI have many different life forms – – – – Yeasts Mycelia Fruiting bodies Spores What do fungi do in soil ? Decompose recalcitrant organic compounds Penetrate residues Stabilize soil structure Nourish fungivores Form symbioses with plant roots and soil fauna Compete with plant pathogens Parasitize plants and soil animals Produce toxins e.g. aflatoxin produced by Aspergillus flavus on peanut Mycorrhizal diversity Ectomycorrhizae AM endomycorrhizae Arbutoid mycorrhizae Ericoid endomycorrhizae Orchid endomycorrhizae Lavelle and Spain (2001) Mycorrhizal Networks: Connecting plants intra- and interspecifically •Many plants are connected underground by mycorrhizal hyphal interconnections. •Mycorrhizal (AM) fungi are not host specific. Illustration by Mark Brundrett Increase nutrient (P) uptake suppress pathogens Mediate plant competition Improve soil structure Glomalin Superglue of the soil ?? PROTOZOA Ciliates • • Largest of the protozoa Move by means of hair-like cilia Amoebae • • Also large Move by means of a temporary foot (pseudopod) Include testate amoebae (with shell-like covering), and naked amoebae • Flagellates • • Smallest of the protozoa Move by means of flagella (1-2) Important bacterivores Soil-Dwelling “Vampires” Vampyrellid amoebae attacking Gaeumannomyces graminis, the fungus that causes “take-all disease” in wheat. NEMATODES Most abundant of the soil fauna Functionally diverse Bacterivores Fungivores Herbivores Predators Omnivores Most are beneficial promoting decomposition and nutrient recycling http://nematode.unl.edu/hdigonic.htm Nematode diversity Bacterivore Fungivore Plant parasitic Predatory Galled root system of tomato infected with root-knot nematode, Meloidogyne sp., compared with noninfected root system Root knot juvenile penetrating a tomato root http://www.agnr.umd.edu/users/nrsl/entm/nematology/images/eis143.jpg Arthropods exoskeleton segmented body jointed legs Arthropod diversity Litter Shredders Fungivores Predators Herbivores Microarthropods EARTHWORMS anecic ~ 35 species of earthworms have been found in Illinois soils ~ half are exotic species from Europe and Asia endogeic epigeic A new earthworm parasite has arrived in IL After earthworms invaded Before earthworms invaded A new earthworm predator has arrived in IL The geographic distribution of Bipalium adventitium in Illinois is unknown. Home gardeners and nursery workers may see these exotic land planarians under boards and stones or crawling on the soil surface after rains. They are up to 2.5 inches long and 1/8 of an inch wide when crawling. They are pale brown or tan with a thin dark brown line running down their back. Scientists would appreciate receiving reports of any observations of these creatures. Contact Ed Zaborski at (217) 265-0330, or zaborski@uiuc.edu by e-mail Epigeic species can be used for vermicomposting Earthworm cocoons http://soils.usda.gov/sqi/soil_quality/soil_biology/images/CE8a-cocoon_LR_small.jpg How do earthworms move through soil ? Loose soil They insert their anterior tip into a pore, extend their setae to anchor most of their body and then exert radial and longitudinal pressure. A cycle of alternating contractions and relaxations thrusts their body forward. Dense soil They invert their pharynx, exude lubricating mucus and suck obstructive soil into their body. How do earthworms eat ? Earthworms eat by pulling food into their mouth with their prostomium (mouth) and pharynx which creates a very strong suction. The food is stored in the crop and then ground up into small digestible pieces in the gizzard. Earthworms need a gizzard because they do not have any teeth. Nutrients are absorbed in the small intestine. Where are the soil organisms ?? Soil organisms are concentrated in HOT SPOTS ! drilosphere detritusphere porosphere aggregatusphere rhizosphere Adapted from Coleman et al. (19??) middens casts Drilosphere Zone of earthworm influence Which burrows were made by an endogeic ? Capowiez et al. 2003) In 1881, Darwin reported that healthy earthworms sometimes leave their burrows at night and embark on a “voyage of discovery” Surface migration is a routine activity aimed at dispersal and finding a mate. Surface migration usually occurs during cloudy moist conditions. Both anecic and endogeic earthworms are active surface migrants. Rhizosphere < 10 % of soil volume Zone of root influence Navigating the rhizosphere Rhizoplane EndoRhizosphere End of the rhizosphere Ecto-Rhizosphere < 10% of soil volume Root free soil > 90% of soil volume (Lavelle and Spain, 2001) Detritusphere surface residue zone Structural rigidity Tortuous, loosely connected and highly constricted porosity The soil matrix presents its inhabitants with many challenges Moisture fluctuations Low quality nutritional resources Microorganisms have very limited ability to move within the soil matrix. As a result, most soil microorganisms are For their prince charmings in a dormant state to arrive ! waiting… Rain Roots There are many types of prince charmings Organic amendments Tillage Soil fauna have a disproportionate impact on nutrient cycling, energy fluxes and plant growth >90% metabolic activity Soil fauna are relatively mobile but have limited digestive ability 3 main types of digestive interactions occur between soil fauna and microflora Soil microflora are relatively immobile but have almost unlimited digestive ability Microbivory NH4 NH4+ + NH4+ NH4+ NH4+ NH4+ Microfauna (e.g. protozoa and nematodes) harness the microflora’s digestive abilities by grazing on them I want some bacteria for lunch ! External rumen digestion The fecal pellets many mesofauna contain Reingestion of fecalofpellets after a few days of shredded, moistened mixed but largely microbial activity greatlyand increases assimilation undigested residues Rumen digestion greatly Soil macrofauna areof enhances utilization ecosystem engineers complex substrates by soil macrofauna Impact of tillage on the soil food web Adapted from Hunt et al. (1986) Soil organic matter fuels the soil food web Active OM energizes biologically mediated processes SOM is a complex mixture Biologically active SOM Living organisms Recent residues Recalcitrant SOM HUMUS Adapted from Magdoff and Weil (2003)