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Home , Fungus, Multicellular organism

Multicellular Eucaryotes most single cells take in and oxygen and get larger size is usually achieved only by being rid of wastes by simple in and out of multicellular the

! cells can’t get too large without breaking in larger there are more ways to solve these problems

better to make lots of small cells than a eg. getting food: , , fluid feeders, few very large cells insectivores, etc multicellularity evolved many separate among eg. getting oxygen: gills, book gills, , tracheae, etc : ! of multicellularity is correlated with an slime explosion of diversity in the record fungi 3. multicellular becomes more resilient

damage doesn’t kill you !must have been a clear advantage

Advantages of Multicellular : !it can often be repaired

1. allows individual cells to become more specialized but there are some tradeoffs: for specific functions: Disadvantages of Multicellular Life: all cells don’t have to do everything 1. individual cells become completely dependent on eg. epithelial cells: cover, protect, support, etc each other eg. muscle cells: movements eg. vascular tissues: transport of and wastes and ! can no longer survive on their own

2. allow much more variation in size & shape

Fungi, Ziser, Lecture Notes, 2011.3 1 Fungi, Ziser, Lecture Notes, 2011.3 2

2. larger organisms need MUCH MORE food and Fungi - General oxygen like the , fungi are a largely invisible ! lots more cells to feed

3. as size increases need more elaborate ways to simplest of three complex multicellular eucaryotic supply each cell with nutrients and remove wastes kingdoms in terms of structure

elaborate systems to collect and distribute nutrients and get rid of wastes no true tissues or organs (as in Plants and Animals)

eg. vascular systems, etc over 100,000 described !est to 1.5M

4. need method of coordination and control to get fungi are extremely abundant in some all parts on the “same page” but not commonly seen:

eg. hormones, nervous systems eg. top 20 cm (8”) of soil contains over 2 tons of (mainly fungi)/acre 5. need some kind of support, especially on land, to counteract gravity (or ~ 2 tons of fungi and bacteria per acre (=5 metric tons per hectare (=2.47 acres)))

6. Dispersal becomes more of an issue eg. a teaspoon full of soil contains several miles of fungal filaments eg. most bacteria and have worldwide distributions most fungi are hidden and inconspicuous and only the relatively few large animals and plants have worldwide “fruiting bodies” are regularly seen: distributions

eg. , , morrels, shelf fungi, etc

some of the largest fruiting bodies:

eg. one species of shelf () weighs 100 lbs (45 kg)

Fungi, Ziser, Lecture Notes, 2011.3 3 Fungi, Ziser, Lecture Notes, 2011.3 4 eg. another shelf fungus in Southern England is 16 feet in The Origin of Fungi circumference and is estimated to weigh ~700 lbs (316 kg) Fungi and Plants are the only kingdoms that are eg. one species of giant can grow to almost 9 feet mainly terrestrial diameter and weigh over 48 lbs (22kg)

fungi appear in fossil record at about the same many are familiar with fungi only due to the and were associated with the first land and economic damage they cause: (some even take it personally): plants

! French Botanist=S Veillardi said fungi “are a cursed tribe, an The appearance of the and Fungi Kingdoms invention of the devil, devised by him to disturb the rest of created by God.” probably occurred before animals moved onto land

In Medieval times fungi were associated with witches, oldest fossil fungi are from ~400 MY ago wizards and magic molecular evidence suggests that fungi emerged ~0.5 - 1 BY ago fairy rings that appeared overnight were said to be the result of fairies dancing in a circle while but no true until ~460 MY ago humans slept molecular evidence indicates some protozoan was the study of fungi was frowned upon by the ancestral to the fungi (and probably animals as )

Church and therefore our knowledge of the at the time () that simple plants began group lagged behind other kingdoms until the producing woody trunks and creating the first 1800’s some of the largest fungi appear in the

fossil record today fungi are known generally as “friends” rather

than foes: ! featureless columns some over 15’ tall

represent an extinct fungus = Prototaxitis providing & and numerous other recycling trash and organic ! probably fed off of microbial crust of bacteria, and covering much of the more barren land of the 1 Million or so suspected species, only a

handful are capable of causing

Fungi, Ziser, Lecture Notes, 2011.3 5 Fungi, Ziser, Lecture Notes, 2011.3 6

Characteristics of Fungi the cells of most fungi have flexible cell walls 1. fungi have colonized all major habitats but are of found mainly in terrestrial ecosystems like , this is a highly indigestible starch: a Nitrogen containing ; similar but some are aquatic to of )

a few are marine, others found in freshwaters most are aerobic

a few inhabit seemingly inhospitable habitats a total lack of O2 prevents growth of most eg. inside glaciers fungi eg. acidic waters -but some can make anaerobically 2. fungi are the most resilient of land eukaryotes but will use oxygen when present

-a few fungi that live in sewage or intestines many fungi are less sensitive to acidic pH’s, higher can also grow anaerobically salt (solute) concentrations, and to colder temperatures than are most bacteria 4. all fungi are nonmotile, they do not even produce motile cells eg. pH! range 2-9; optimum = 5.6 5. most, not all, fungi are multicellular eg. thrive in a wide range of temperatures

some can thrive in freezing temps yeasts = general term for unicellular fungi

!common in refrigerators molds = general term for multicellular fungi eg. thrive in high osmotic pressures !high sugar or salt concentrations, jellies, jams multicellular fungi remain much simpler in structure than higher plants and animals 3. cells of fungi are eukaryotic (have nucleus & various ; mitochondria, , etc) ! no true tissues or organs

cells are enclosed within cell walls

Fungi, Ziser, Lecture Notes, 2011.3 7 Fungi, Ziser, Lecture Notes, 2011.3 8 except for yeasts, bodies consists of filaments when growing is limited, the fungus infuses called hyphae the food with hyphae and assumes the shape of its food. in most the hyphae consist of chains of individual cells a can get quite large:

but the cell walls do not completely separate ! fungi may be ’s oldest and largest adjacent cells organisms on earth

! essentially, all cells in hyphae are interconnected: eg. fungus growing beneath coniferous in Michigan acts as one long single celled filament 37 acres 11 tons of hyphae the hyphae form an interwoven mat = mycelium 1500 old ( growers call it “”) of individual feeding filaments (=hyphae) extending into eg. ostoyae (Washington State) 2.5 square miles – covers three counties their preferred food supply probably 1000’s of years old probably 100’s of tons mycelium represents the individual organism eg. an individual crustose (a between a fungus and algae is estimated to be up to 3,700 years hyphae grow very quickly old

a single hyphal filament that grows and branches can grow 6. fungi are like animals over a half mile of hyphae in 24 hrs; in 2 days, 100’s of miles a. most are saprobes () that feed the filamentous structure provides an extensive on dead surface area for absorption of nutrients: particularly good at digesting plant compounds the vegetative portion of most fungi are often such as cellulose and lignin hidden in the food they are eating ! very common on dead trees and rotting logs

fungi are organisms with no clear shape: but same substances cause fungal attacks on clothing, always growing and changing paper,shoes, insulation, lumber

Fungi, Ziser, Lecture Notes, 2011.3 9 Fungi, Ziser, Lecture Notes, 2011.3 10

also thrive on a great variety of other organic most of the mass of the fungus grows into the food it eats compounds and absorbs nutrients

2. rings and traps asbestos – that predatory soil fungi use to capture prey jet fuel

rubber -some have adhesive ends leather th -some rings can constrict or swell in a 1/10 of a second

-some prefer certain species as prey b. some fungi are parasites -once prey are “caught” hyphae penetrate the prey and digest and dissolve it then absorb nutrients eat living organic matter eg. hyphae of the common mushroom,

Pleurotus ostreatus some can switch between saprophytic and parasitic

3. parasitic hyphae (haustoria) some are parasitic in plants and animals, –hyphal tips of parasitic fungi that invade cells of including humans -secrete to dissolved their way through cell walls and cell membranes ! about 300 known animal 4. reproductive hyphae ! most important cause of agricultural plant diseases - may form large visible “fruiting bodies” that produce sometimes get established in areas making it difficult to get agricultural to grow – usually the most visible part of the fungus - reproductive hyphae produce either asexual or sexual spores c. a few are even predators

7. most fungi reproduce by sexual and asexual spores

yeasts reproduce asexually by

8. fungi lack true tissues but hyphae have become specialized for a variety of functions:

1. feeding hyphae (=vegetative hyphae)

Fungi, Ziser, Lecture Notes, 2011.3 11 Fungi, Ziser, Lecture Notes, 2011.3 12 of Fungi fungi play important role in recycling organic nutrients in many ecosystems fungi produce a variety of distinctive chemicals particularly good at digesting plant compounds 1. digestive enzymes such as cellulose and lignin

2. ! very common on dead trees and rotting logs 3. bioluminescent chemicals especially important in recycling dead many of the chemicals that fungi routinely produce cannot yet be trees synthesized in the laboratory

1. Digestive Enzymes !they are some of the few organisms that can digest cellulose all fungi are heterotrophs along with some bacteria and protists

fungi are found wherever organic matter is but fungi make no distinction between available a rotting tree, a railroad tie, cloth, paint, leather, insulation, etc but grow best in dark, moist habitats they also thrive on a great variety of other most fungi are saprobes (=decomposers) organic compounds

!eat dead organic matter asbestos rubber leather some are parasitic plastic wax fungi secrete a wide variety of enzymes to glue

digest organic molecules: also attack printed circuits, coatings on

camera lenses, kerosene fungi can decompose almost any organic molecule eg. the “kerosene fungus” eats jet fuel

Fungi, Ziser, Lecture Notes, 2011.3 13 Fungi, Ziser, Lecture Notes, 2011.3 14

also grow on foods, grains, bread, , the fungus concentrates its energy and resources by vegetables including those in cans and in constantly adding more area for absorption into new areas freezers virtually all growth is in the form of elongation of the tips of ! cause the hyphae

a single fungus can grow ~ 1/2 mile of hyphae per day while some fungi can grow on an extremely wide variety of foods others are so specialized that once original food supply is depleted, the they grow on only a single hyphae explore areas further distant leaving an area exhausted of food eg. one fungus only grows on impregnated surrounded by a growing ring of hyphae

eg. a specific species of pollen, or feathers ! “fairy rings” on lawns

eg. of specific tree species, only on certain insect species, etc ! ringworm on skin

fungi are absorptive heterotrophs: 2. Mycotoxins

!they must eat organic foods for nutrients while plants and animals have a protective “skin” and energy covering mushrooms and fungi do not

they reverse the animal technique of taking in ! leaves them much more susceptible to food and then digesting it bacteria and

they secrete enzymes to digest food outside at same time that they are secreting enzymes their bodies, then absorb it they are also secreting mycotoxins

fungi make up for lack of mobility by growing mycotoxins = chemicals produced by fungi to rapidly “protect” their food source from bacteria and other fungi

inhibit growth of other microbes Fungi, Ziser, Lecture Notes, 2011.3 15 Fungi, Ziser, Lecture Notes, 2011.3 16 Environmental Resistance ! possibly to discourage for food !many fungi can grow at temperatures below freezing

some of these we use as other fungi do not die, they stop growing

some of these mycotoxins, in humans, are some have been exposed to temperatures below 400º and they remain viable however most fungi are killed at 100º C 3. ! heat treatment is used to kill fungi on seeds some fungi are bioluminescence !fungal spores are also quite resistant to high >40 fungal species glow in the dark temperatures and

in some cases may improve reproductive success !in some, when environmental conditions are unfavorable or food supply runs out, hyphae may eg. bioluminescent fruiting bodies aggregate into a roughly spherical body of olearius, stypticus, illuminens = scerotium

perhaps: attracts insects that help to disperse spores outside cells – thick pigmented walls form protective shell may attract animals such as protect agains UV which eat fungus and excrete undigestible spores when favorable conditions return the eg. bioluminescent mycelia may germinate into hyphae again or produce a fruiting body unsure of function; apparently not always reproductive =foxfire of lore : have been used to mark trails at night

used by troops to identify friend from foe most fungi reproduce both asexually and sexually by producing spores to brighten hair of young maidens on tropical isles reproduce asexually most of the time

Fungi, Ziser, Lecture Notes, 2011.3 17 Fungi, Ziser, Lecture Notes, 2011.3 18

reproduce sexually when conditions change and become less puffballs, etc favorable

Asexual Reproduction: ! there are numerous different “ types”(genders) in fungi most fungal reproduction involves the production (not called male and female) of asexual spores [some fungi can have up to 78,000 different asexual spores are produced on the modified mating types or “” in a single species] tips of reproductive hyphae also, unlike plants and animals they do not form microscopic; no large fruiting body visible

only visible as “colored, fuzzy molds” germinates directly into new hyphae

whenever conditions are favorable fungi a few fungi reproduce parthogenetically themselves by producing millions of asexual spores from specialized fruiting bodies Number of Spores

yeasts reproduce asexually by budding the number of spores produced is often enormous:

eg. a common wood rot fungus produces 30 billion : spores/day or 350,000/second up to 6 months each

sexual spores are generally much less commonly eg. corn ! 25 billion spores/ ear of corn

produced than asexual spores eg. ! 10 billion spores/ acre (moderate )

~1/3rd of all fungi reproduce in more than one way eg. some club fungi: ! 80 Million spores/hour for several days

eg. sexual and asexual Dispersal of Spores

sexual spores are produced on large fruiting fungal spores are dispersed by: bodies

a. visible to naked eye as mushrooms, Fungi, Ziser, Lecture Notes, 2011.3 19 Fungi, Ziser, Lecture Notes, 2011.3 20

b. projectile dispersal fruiting bodies have gills, long tubes or teeth from which the spores are released c. eg. shaggy mane () d. insects produces its spores on long deep gills

e. as gills on outer rim of mushroom release their spores, the fungus secretes enzymes to digest this area a. Wind and the next area for spore release

Most sexual and asexual spores are dispersed by if a fruiting body is tilted or falls over, spores wind cannot be dispersed

many of the more destructive plant disease fungi some shelf fungi will regrow a new fruiting produce spores dispersed by wind body in the correct orientation

eg. ; a shelf fungus smaller spores will remain aloft longer if tree or log falls it will grow new shelves to eg. dish opened on 1st floor in office bldg realign tubes for dispersal

in 5 minutes some spores were on 4th floor b. Projectile Dispersal

th in 10 minutes they were on 4 floor at densities of some spores get help getting into the wind: 2 1000’s/yd eg. many ascomycetes have cup-like fruiting bodies also spores are usually resistant to adverse conditions ! spore must get out of cup into wind for dispersal

spores are inside sac ! can survive long periods in transit the exposed end of sac has hole in it or is hinged

most spores must have clear access and use changes in water pressure within causes tip to give gravity to fall out of fruiting body way

Fungi, Ziser, Lecture Notes, 2011.3 21 Fungi, Ziser, Lecture Notes, 2011.3 22

slime lining “explodes” and hurtles spore packets and slime spores are shot out through the end out of cup ! 27 ft/sec thick of spores can shoot several inches ! to 18 ft away into the air you can hear them “go off” then are caught by wind and dispersed c. Rainfall eg. (Ascomycete)

eg. some wood disease causing fungi grows on horse dung (rich in undigested cellulose,nitrogen,

etc.) produces fruiting bodies that look like small flasks under

bark of tree spores were eated with plants and are released in dung

when bark is moistened by rain spores ooze out of flask in as soon as dung is released, Pilobolus absorbs water and sticky gooey mixture sends up a small fruiting body (2-4 cm)

like toothpaste out of tube sporeheads on fruiting body aim toward light

eg. birds nest fungus ( Basidiomycete) fruiting body slowly dries

2 produces packets of spores inside cuplike “nest” when internal pressure exceeds 700 kg/cm sporeheads up to 1/3rd “ diameter pop out of and land several meters away in

grazeable grass each cup contains 4-10 packets

eg. water dropping on nest during rain causes packets to

splash up the sides of “nest” grows on decaying wood

some have teather like filaments with sticky ends that help common in wood chips used for mulch, esp NE US to fasten packet to nearby objects

spores produced inside cup packets and dry to release spores

cup is covered with slimy substance d. water dispersal on a rainy day when spores are ready slime absorbs water and swells

pressure builds

Fungi, Ziser, Lecture Notes, 2011.3 23 Fungi, Ziser, Lecture Notes, 2011.3 24 spores of often have “appendages” infects rock cress (mustard) in Colorado Mtns that allow them to float longer in the water once infected the fungus makes a nectar rich rosette resembling a flowerhead and therefore enhance their dispersal it attracts “would be” pollinators to spread spores e. Insect Dispersal iv. bioluminescence:

many spores dispersed by insects are sticky eg. some fungi apparently use bioluminescence to attract insects to spores

must have some way to attract the insect to the e. Mammals fungus for dispersal eg. Morels & i. nectar: many of the fruiting bodies have a strong smell and flavor

! excavated and eaten by animals ( & eg. and Rusts rabbits) (including humans)

plant parasites ! dispersal need to get from host to host

deposit spores in sweet nectar eg. melanosporum (Black ) insects are attracted to the nectar and disperse

spores contains alpha-androsterol a also found in breath of

male pigs (and male and female humans) ii. smell:

eg. stinkhorns

produce spores in a slimy, stinky mass resembles rotting stench attracts flies and insects to disperse spores 100’s of flies descend and can remove all spores in a few hours

iii. :

eg. yellow rust fungus

mimics a Fungi, Ziser, Lecture Notes, 2011.3 25 Fungi, Ziser, Lecture Notes, 2011.3 26

Ecological Roles of Fungi lignin appeared millions of years after woody plants evolved 1. fungi play important role in recycling organic the earliest fungi (before woody plants evolved) were nutrients in terrestrial ecosystems not yet able to digest lignin

eg. In forest soils other than plant the fungi are the when woody trees and shrubs evolved, it took fungi dominant life forms. several million years of to adapt to this new food source and be able to produce fungal is 90% of the total organic matter enzymes that could digest this lignin

All the rest together including nematodes, , This lag of several million years in fungi ability to digest bacteria, , , snails, insects, insect lignin explains why early forests of woody plants larvae, etc weigh the other 10%. were not decomposed

eg. In agricultural soils the biomass of fungi is equal to the !and become massive coal deposits (250-300MY) weight of all other living organisms combined (1250 kg per

hectare). No major coal deposits have formed since the even though there were many more fungi can decompose almost any organic woody plants since then molecule

especially important in recycling dead trees 2. Fungi also play a vital role in and energy cycling in freshwater ecosystems !they are some of the few organisms In streams and rivers most of the energy comes from wind blown that can digest cellulose leaves and plant debris rather than from by plants or algae along with some bacteria and protists The plant debris is high in but low in nitrogen (high !only fungi can digest lignin C/N ratio)

(an important component of wood) Aquatic fungi colonize the leaves and lower the C:N and make them more palatable (nutritionally useful) to feeding arthropods (such as Gammarus) that are at the base of the foodchain.

3. Role in Carbon Cycle

Fungi, Ziser, Lecture Notes, 2011.3 27 Fungi, Ziser, Lecture Notes, 2011.3 28 85 billion tons of carbon is returned annually to the atmosphere ! fertilized fields reduce diversity of fungi as by decay of cellulose and lignified cellulose. Most of this is by fungi. ! polluted waters on lands reduce fungal diversity 4. recent discoveries indicate that a few fungi might actually be ! atmospheric pollution also decreases diversity

they don’t do photosynthesis and extract energy from

instead they are able to extract energy from radioactive minerals using the to absorb the

and use that energy to make sugars

! Fungi at Chernobyl may be a new kind of ??

1986: 200 tonnes of radioactive fuel melted inside reactor

1999 a robot sent into the reactor returned with a sample of a very black fungus

dark pigment was melanin (skin pigment)

later found similar melanized fungi growing in cooling ponds of other nuclear reactors

melanin protects us from various forms of radiation

here melanin seems able to capture energy from radiation and transform it to useable energy

grows much faster in radiation than other fungi with or without radiation

somehow the fungus seems to harvest energy from through the melanin;

but still needs further investigation

Today, fungi are affected by habitat loss and pollution

Fungi, Ziser, Lecture Notes, 2011.3 29 Fungi, Ziser, Lecture Notes, 2011.3 30

Symbiotic Fungi Lichens:

Fungi are noted for their numerous symbioses with a symbiotic association between a fungus and a other organisms photosynthetic partner

! they commonly form symbioses with ~>20,000 “species”; generally known by the binomial name of the fungus partner a. bacteria (b-g) or algae lichens the symbioses produce a body form different from that of either the fungus or the alga b. plants mycorrhizae = c. animals ‘fungus farms’ a thallus can have any of three main shapes:

crustose ! crust like spreading growth

foliose ! thin crustlike sheet but with raised edges

fruiticose ! shrublike branching growing up from substrate

the thallus is mainly made of fungal cells (>90%)

!fungi generally produce the “housing”

the photosynthetic organism is usually a green alga and usually comprise only ~ 7% of the lichen

sometimes it is a blue-green bacterium or both

the algae (but not the fungus) can also be found free living on its own

Fungi, Ziser, Lecture Notes, 2011.3 31 Fungi, Ziser, Lecture Notes, 2011.3 32 the algae produces sugars from photosynthesis eg. dry desert, cold frozen north

bg bacteria can also fix nitrogen and provide other organic lichens absorb minerals mainly from the air and nutrients for the fungus rainwater

most (up to 90%) of the sugars produced by the lichens produce some unique secondary chemicals algal cells is transferred to the fungus !prevent other lichens from growing nearby lichens vary greatly in size: microscopic to Km’s ! protect lichen from UV radiation grow very slowly ! some are chemicals that dissolve and weather the rock to

release minerals that can be absorbed by lichen eg. many crustose lichens grow only mm/century

some lichens are 1000’s of years old, may be some of the lichens reproduce mainly asexually by fragmentation oldest living lifeforms on earth ! bits of lichen break off lichens are the dominant organisms over 8% of the earth’s land surface What Kind of Symbiosis? lichens can grow virtually anywhere: lichens were originally considered an excellent example of from deserts to tropical

in the Anarctic, in freshwater streams, in the rocky intertidal but we now know the relationship is parasitic: areas of seacoasts

on fences, rocks, power lines, houses, gravestones, even on ! the fungus produces haustoria that actually turtle shells penetrate the algal cell they can tolerate extreme temperatures and lack of ! the fungus absorbs up to 90% of the sugars moisture produced by algal photosynthesis

often abundant in habitats too harsh for most ! if bg bacteria, the fungus also absorbs up to 95% of the fixed other forms of life nitrogen () made by the bacterium

! algal (or bacterial) cells slowly die Fungi, Ziser, Lecture Notes, 2011.3 33 Fungi, Ziser, Lecture Notes, 2011.3 34

eg. grouse and wild turkey eat lichen. alga actually grows better alone than in symbiosis with the fungus eg. Warblers, golden plovers and Vireos use lichens in their nests ! the fungus is never found growing on its own eg. The -headed weaver in Madagascar makes its nests solely from lichen in the . ! without the alga, the fungus requires numerous organic compounds to grow eg. antelope, , elk and small mammals also utilize lichen.

Ecological Importance of Lichens Human Uses of Lichens 1. lichens are considered : 1. air pollution indicators often first organisms to colonize an area lichens absorb minerals mainly from the air and rainwater !facilitate weathering process and soil formation lichens cannot excrete elements that they absorb

2. in extreme habitats lichens may be the ! some are very sensitive to air pollution (esp SO4, Ozone, Flourides) predominant primary producer and the base of the lichens thus require clean air to thrive

eg. lichens are especially abundant in the arctic tundra lichen diversity is used to map the spread of air pollution around large cities over 350 sp lichens only 2 sp flowering plants 2. As food

“reindeer ” is an important food source for caribou and reindeer in the arctic Lichens are eaten by people in North America, Europe, Asia, and Africa note: “reindeer moss” is not a moss but a lichen Often lichens are merely famine foods eaten in 3. Food & nesting materials for wildlife in many times of dire needs, but in some cultures terrestrial ecosystems lichens are a staple food or even a delicacy. eg. In the past Iceland moss was an important human Fungi, Ziser, Lecture Notes, 2011.3 35 Fungi, Ziser, Lecture Notes, 2011.3 36 food in northern Europe and Scandinavia, and 6. Other human uses of lichens was cooked in many different ways, such as bread, porridge, pudding, soup, or salad Lichens have been and are still being used for many other purposes

Alcohol production (for fermentable , as eg. Northern peoples in North America and catalysts, and/or as flavour/preservatives) traditionally eat the partially digested lichen after they remove it from the of caribou that Cosmetics (for hair, and/or sweet smelling powders) have been killed. It is often called 'stomach Perfumes icecream'. Decorations (including costumes and artwork) eg. is a lichen that has frequently been used Fibre (clothing, housing, , sanitation) as an emergency food in North America. Animal feed (both fodder and ) eg. “manna” of biblical fame was probably a local lichen in the area Industrial purposes (production of acid, carbohydrate)

3. some provide yellow, red and purple pigments for

eg. wolf lichen

4. litmus is a lichen pigment used as a pH indicator

5. Pharmaceutical products from lichen extracts:

It has been estimated that 50% of all lichen species have properties.

eg. usnic acid used in antifungal and antibacterial products

eg. reindeer moss ! expectorant, taken for colds and as a laxatve

eg. ! to treat sores

eg. pixie cup lichen ! antibiotic for TB

Fungi, Ziser, Lecture Notes, 2011.3 37 Fungi, Ziser, Lecture Notes, 2011.3 38

Mycorrhizae mycorrhizae are also important in water uptake and storage (means “fungus ”) ! commerical plants need less irrigation symbiosis of fungus with plant root cells ! don’t suffer as much under water stress as plants grown without mycorrhizae the symbiosis was 1st discovered in mid 1800’s earliest fossil records of land plants contain most “mushrooms” found in the are the fruiting evidence of mycorrhizae (>400MY) bodies of the fungi in these mycorrhizal relationships ! played major role in evolutionary step that brought aquatic plants to land: Each forest tree has hundreds of thousands of miles of hyphae extending from its roots greater area allowed plants to leave aquatic environment and exploit relatively harsh soil over 90% of all vascular plants (most perennial environment plants) have mycorrhizae there are some plants (eg. most orchids, some nearly all commercially important plants have trees and shrubs) that have no root at all micorrhizal associations and depend almost completely on mycorrhizae for sustenance

The symbiosis is a form of mutualism in some their seeds won’t even germinate unless the fungal partner is with them Benefits to Plant: 2. mycorrhizae release “powerful chemicals” into soil 1. greatly extend the absorptive ability of roots to dissolve hard to capture (ie. tightly bound) nutrients such as phosphorus and iron ! increase the absorptive area 10-1000 x’s ! plants grown without mycorrhizae often require high mycorrhizae are able to absorb and transfer all levels of fertilizer to grow as well 15 major and micronutrients needed for plant growth.

Fungi, Ziser, Lecture Notes, 2011.3 39 Fungi, Ziser, Lecture Notes, 2011.3 40 3. mycorrhizae are able to excrete specific antibiotics to attack plant pathogens entering soil aeration & soil structure root zone mycorrhizae produce humic cmpds and organic “glues” (=glomalins) that promote aeration, root growth and water also acts as a physical barrier to pathogens movement

they create a ‘sock-like’ covering of tightly interwoven soil water uptake hyphae around root hairs

Benefits to Fungus: many common practices can degrade mycorrhizal development: 1. plant provides refuge from hostile conditions tilling fertilization

removal of topsoil !allows fungi to withstand drought and cold erosion

road and home construction 2. provide major energy source to fungus in return fumigation for its “services” invasive plants bare soils !up to 25% of food made by plant is “given” to fungus nursery grown plants are often deficient in Ecological and Economic Importance of mycorrhizae Mycorrhizae ! need more watering and fertilizer and very important in natural ecosystems and in use sterile soils agriculture and in industry ! plants poorly adapted for transplanting to more most plants usually grow poorly when deprived of natural areas their mycorrhizal symbiont but: can use inoculum when transplanting mycorrhizae have important effects on: in some cases plant diversity of inoculated & nutrient cycling replanted plots is ~2x’s that of uninoculated sites extract nutrients from poor soil Fungi, Ziser, Lecture Notes, 2011.3 41 Fungi, Ziser, Lecture Notes, 2011.3 42

the earlier the inoculation the better Animal “Fungus Farms”

eg. (leafcutter ants)

mounds up to 18 ft underground

the ants maintain active fungal gardens to feed larvae and adult in the

!they cultivate fungi within their mounds

!they feed the fungi bits of leaves, bark and other plant material

!they out debris that might spoil their garden

!have wheelbarrow-like depression on their backs for carrying spores

fungus is transferred from colony to colony by ants

free-living cultures of fungus have never been found

fungus needs ants to provide them with food

in Central and South America leaf cutter ants are the dominant herbivores in food chains

! consume 17% of total leaf production

eg. African (Microtermitinae)

build chimneys up to 30 ft tall of feces and mycelia

Fungi, Ziser, Lecture Notes, 2011.3 43 Fungi, Ziser, Lecture Notes, 2011.3 44 Human Impacts of Fungi eat mostly plant material but don’t have cellulose digesting microorganisms living in their gut Former uses of fungi:

bring it home and eat it before entering in ancient times fungi were used for such as mound putting on wound to help it heal

! related to antibacterial properties of many fungi then go in and defecate puffball spores encourgage clotting of wounds use their feces to grow fungi () fungi were used to make the spark for muskets and as tinder for fires all stages of readily eat the fungus 1. all crops, garden plants trees and grasses fungus never forms mushrooms in mound depend on mycorrhizae to help them grow and produce commercial products but when rain is imminent 2. Fungi are eaten directly as Food termites carry bits of mycelia out of mound

spread them around the ground generally an excellent source of , minerals and fiber with very small quantities of fats several days after rain mushrooms appear and produce spores over 200,000 tonnes fungi are eaten each year

the termites reappear and collect the genetically mixed hyphae and take them back into the nest 200 species of edible mushrooms

abandoned mounds produce crops of mushrooms some are cultivated commercially for several years after termites leave eg. brunesscens (supermarket fungus) ! African people consider them a rare grown in large caverns in Peru delicacy has been thoroughly domesticated

Many other mushrooms and fungi are edible Fungi, Ziser, Lecture Notes, 2011.3 45 Fungi, Ziser, Lecture Notes, 2011.3 46

and commonly used in soups and stews as natural truffles are collected using trained well as main dishes pigs or dogs or following “truffle flies” to source

eg. Woodear Mushrooms (=tree ears, black fungus) truffles are some of the most highly priced “produce” in the market eg. Mushrooms (=oak mushroom): they have been sought out for grows on oak logs or cultivated on oak chips but: some larger fungi can accumulate high eg. Oyster Mushrooms ( ostreatus): levels of metals and radioactive isotopes grow on dead and decaying hardwoods or can be (=biomagnification) cultivated on sterilized straw ! don’t eat fungi along highways or in heavily eg. elephant ear fungus polluted areas used in hot and sour soup 3. Fungi used to manufacture foods & sauces thought by some to prevent and retard heart disease (not proven) eg.

eg. Morels & Truffles yeasts are used commercially for producing considered gourmet fair bakery products and alcoholic beverages !fermentation produces many of the fruiting bodies have a strong smell CO2 and flavor alcohols France grows specific species of oak trees to act as hosts for truffles Yeasts are single celled fungi. Baker’s yeast, cerevisiae, ferments the sugars in a each truffle takes 7 years to mature mixture of dough to produce carbon dioxide which causes the dough to “rise” (alcohol is vaporized) eg. (Black Truffle) eg. so far has been impossible to grow commercially The first step in making most cheeses is to prepare a curd by adding bacteria and rennin or bacterial enzymes to milk.

Fungi, Ziser, Lecture Notes, 2011.3 47 Fungi, Ziser, Lecture Notes, 2011.3 48 The bacteria sour the milk and enzymes coagulate the milk A. oryzae , casein, to produce a soft ‘curd’ to make Saccharomyces rouxii and a liquid ‘whey’ which is a waste product. versitilis

The amount of whey removed determines the hardness of The starch degrading enzymes produced by the molds the cheese. eg. soft cheeses the whey is simply produces a sugar that is then fermented by the bacteria. drained away. Harder cheeses are heated and The entire process takes about a year. pressurized to remove additional whey from the mixture. 4. Beverages

After this separation, most cheeses are ripened with eg. Coffee inoculations of various species of bacteria and/or fungi. After coffee beans are picked they are soaked in water containing natural cultures of Erwinia dissolvens and Soft Cheeses Saccharomyces spp. to loosen the berry skins before

roasting. Some fermentation occurs which is believed to Brie, Camembert: The curd is ripened (cured) by a produce some of the unique flavors of various varieties of variety of microorganisms including Streptococcus coffee. lactis, S. cremoris, Penecillium camemberti and P.

candidum. eg. Cocoa

In this case the ripening process is aerobic and the Microbial fermentation by and cultures are inoculated onto the surface of the cheese spp. is used to help remove the cocoa beans from the and extend hyphae throughout. covering them in the pod. The products of this

fermentation contribute to the flavor of the cocoa. : unique flavors and smells produced partly

by Penecillium roquefortii eg. alcoholic beverages

some of the finest French cheeses are distinguished by strains that produce mainly the smell of isobutyric acid produced by fungi (also alcohols on fermentation: helps to produce the characteristic odor of vomit)

: sugars ! ethyl alcohol eg.

Soy sauce is produced by the fermentation of roasted soy beer: barley ! CO2 and ethyl alcohol beans and wheat using a mixture of various bacteria and fungal species including: most of the flavors of wine arise from the direct activity of the yeast rather than the flavor of the bacteria: Pectococcus halophilus grape delbrueckii fungi: soyae 5. Pharmaceuticals

Fungi, Ziser, Lecture Notes, 2011.3 49 Fungi, Ziser, Lecture Notes, 2011.3 50

from and citrinum. fungi produce many and to 6. Industrial Chemicals & Products from fungi compete with bacteria for space and food

eg. (Saccharomyces cerevisiae) ! used extensively for such chemicals often have certain effects on the ethanol production for beverages and biofuels and are often a primary source eg. () ! used in some toothpastes; for medical applications and as digestive aid

eg. Antibiotics: eg. () ! All the citric acid used in soft drinks, candies, artificial lemon juice, baked goods etc. Penicillium genus is source of first antibiotic is produced industrially by fungus fermentation. eg. Fumaric Acid ( arhizus) ! used in food production 1928, Alexander Fleming accidentally discovered its for flavoring and as a preservative antibiotic properties eg. Beano© produced by from Aspergillus niger (=Cephalosporium) produces an : alpha-d-galactosidase suppresses methane production in human digestive tract by Penicillium griseofulvin eg Penicillium bilagi eg. ergot. some drugs are produced from ergot that: makes soluble and more easily absorbed by crop plants induce labor stop uterine bleeding eg. takadiastase (Aspergillus oryzae) ! starch and protein treat high blood pressure splitting enzyme used as a digestive aid relieve migraines cancer treatments eg. some shelf fungi are being used to make specialty paper treat hepatitis B infections eg. many countries use fungal dyes to color yarn eg. cyclosporin over 70 different colors derived from fungi one of least toxic, most effective immunosuppressive drugs known 7. Mycotoxins from inflatum(?deutero?)

eg. poisons produced by some species of fungi that statins are the most effective cholesterol lowering drugs in can cause , sickness or if use today they inhibit enzymes needed to make cholesterol taken in large amounts Fungi, Ziser, Lecture Notes, 2011.3 51 Fungi, Ziser, Lecture Notes, 2011.3 52

eg. poisonous mushrooms produces a structure called ergot.

1st description appeared in Middle Ages about 70 species are poisonous (=toadstools)

Eating ergot can cause nervous spasms, convulsions edible and poisonous can look very similar; and psychotic delusions, may even be in same genus

ravaged Europe in middle ages no simple way to distinguish them

in 994 France >40,000 people died from ergot eg. yellow stainer () poisoning

the most commonly ingested poisonous mushroom caused outbreaks of madness attributed to

possession looks very similar to commonly eaten species

in 1670’s a high death rate for travelers on religious white or off white color; damage produces a yellow pilgrimages was related to eating bread made stain on mushroom; smells somewhat like iodine from contaminated grain

if eaten may cause abdominal cramps, nausea, ! sometimes fingers, toes, arms and legs vomiting or diarrhoea became dry and black with and

sometimes fell off eg. (destroying angel, death cap)

! believed limbs were consumed by the one of most poisonous “holy fire” or “St. Anthony’s Fire” while

on pilgrimages is not neutralized by cooking

eg. one bite is fatal

many are aware and concerned about the dangers of onset of symptoms from 3 to 36 hours after ingesting ingesting synthetic additives or in their

food. death usually from liver or kidney damage

Yet few are aware that we consume over eg. ergot (causes ) 10,000 x’s more of the “natural” poisons in the

foods we eat than the artificial poisons we are best known most worried about

caused by Claviceps that infects the of and one of the better known “natural” toxins is other grains

Fungi, Ziser, Lecture Notes, 2011.3 53 Fungi, Ziser, Lecture Notes, 2011.3 54

produced by species of Aspergillus which grows on , rice, grains, beans, milk eg. most (186) contain psychoactive

causes liver damage and cancer often referred to as “magic mushrooms”

been reported in all parts of the world eg. liberty cap semilanceata

most prevalent in areas of poverty and starvation mimic effects of LSD but 200 times less potent

eg. half of all in Africa south of Sahara mimics effects of (neurotransmitter) are liver tumors ~ half of these are found in Mexico same area also has high incidence of childhood liver cirrosis ! still used in religious ceremonies

correlated with report that 40% of foods eg Psilocybes mexicana screened there contained measurable amounts of aflatoxin some are being studied to reduce anxiety about dying in terminally ill patients including breast milk eg. others contain a different set of such as eg. one such destroyed whole villages muscimol in and Siberia when famine forced villagers to eat moldy grain eg. red-and-white spitted fly () one of main exposure sources of aflatoxin in US is butter used as an intoxicant in Siberia

[organic peanut butter contains much higher eg. some contain alkaloid concentrations than commercial brands. Since its not processed and sterilized as quickly, the fungi get a longer time to grow and produce the toxin] LSD is derived from these

eg. hallucinogens 8. Allergens

ingestion of some mushrooms causes We breathe in fungal spores every day intoxication and sometimes hallucinations clean air may contain as many as 10,000 fungal spores per cubic metre about 200 species of mushrooms are known to be hallucinogenic your pillow alone may harbor up to 1 million spores Fungi, Ziser, Lecture Notes, 2011.3 55 Fungi, Ziser, Lecture Notes, 2011.3 56 Also fungi grow very nicely in and around houses. esp in basements and bathrooms some also infect mucous membranes of mouth, etc

were much more common before soap was regularly used can become a significant component of house dust and

a major problem for those with mould . easily spread

Those who are prone to allergies can become eg. Mycoses sensitized to spores from any of a large number of fungal species. internal infections are called mycoses

some of the most important human allergens & include Penicillium, Aspergillus, and . some cause respiratory infections with flu-like symptoms

fungal infections, particularly internal mycoses are 9. Human Pathogens sometimes difficult to treat and may be lethal in immune compromised patients most human fungal infections of the skin are rarely fatal but some can become disabling or drugs to treat these infections because, unlike bacterial diseases, the chemistry of the fungus is much more disfiguring. similar to our biology ( vs )

a few can enter the blood and become systemic, of the major kingdoms of life, fungi are more like animals than any other kingdom sometimes causing fatal brain infections eg. eg. fungal skin infections (=) Bausch & Lomb recently (2006) recalled contact lenses that a fairly large number of fungi invade skin hair and nails were infected with this fungus causing severe eye infection in some yeasts (eg Candida) and some molds (eg. ) related to poor hygiene with contact lenses especially in areas of body that tend to be damp 10. Food Spoilage cause: ringworm, thrush athlete’s foot and rd “yeast infections” >1/3 of all food harvested worldwide is destroyed once established may be difficult to treat or may persist for by fungi and insects years Fungi, Ziser, Lecture Notes, 2011.3 57 Fungi, Ziser, Lecture Notes, 2011.3 58

therefore can grow in dry locations and along rocks, over $50 B/yr of stored crops are lost mortar, wires and cables until it finds wood

all plant fibers are subject to fungal attack most of the blue-green, pink or brown molds that cause food to spoil are ascomycetes eg. during WWII clothing tents, ropes, etc were under constant attack from fungi, especially in tropical areas eg: Rhizopus (=black bread mold)(zygomycete) in addition to lumber & plant fibers, fungi destroy is an occasional household many other manufactured products:

11. damage to wood & manufactured goods paint leather several 1000 species of fungi feed exclusively on insulation asbestos wood jet fuel rubber wood rot fungi cause millions of dollars of loss printed circuits each camera lenses

year. 12. Plant and Animal Pathogens & Diseases

two main kinds: virtually all living plants & animals are subject to wet rot attacks by fungi dry rot Plants a. wet rot

only attacks wood with a high moisture content fungi are among the most common plant needs damp conditions pathogens rots outbuildings, wet cellars termites are attracted to wood that already has fungal plant diseases cause millions of dollars each damage from wet rot fungi year to crops and produce

b. dry rot fungus; sp. probably originated in the Himalayas and has now plant diseases include smuts and rusts spread throughout the world attack corn, wheat, oats, other grains can produce its own water by as it grows Fungi, Ziser, Lecture Notes, 2011.3 59 Fungi, Ziser, Lecture Notes, 2011.3 60 hard to treat since no single fungicide will kill all the fungus attacks trees through breaks in the bark fungal pathogens and grows in the sapwood

today the species is almost extinct eg. ( fagacearum)

however the American Foundation is working attacks several species of oak trees mainly in midwest to develop blight resistant strains and hope to

“resurrect” the species again in central Texas mainly occurs in older live oaks

also: symptoms quickly appear, usually in spring, and trees dutch disease, die within a couple of months ,

some powdery spread by root grafts and bark scab

brown rot no cure, only preventive trenching can limit spread

Animals (nonhuman) eg. Chestnut Blight ( parasitica)

eg. White nose disease in 100 years ago, American Chestnut tree was the

dominant tree in some ecosystems mysterious fungal disease devastating populations

in NE US. !25% of all trees in east US were American

Chestnuts (~ 9 Million acres) has killed 400,000 bats in last 3 years (2009)

! one of the most common trees used to has recently (2009) been found in Europe line streets and avenues

eg. common housefly fungus (Empusa sp.) mature tree >100 ft tall, some dia 10 ft

most attacks noticed in fall when flies become sluggish take an entire RR car to haul wood from 1 tree

once infected fly becomes more and more sluggish 1904 chestnut blight was introduced into N America

from Asia (bioinvasion) soon can’t fly, can only buzz wings

apparently spread by wind, insects, birds and within a few days the fly dies mammals

when the fly dies, reproductive hyphae puncture its American trees lacked the resistance that Asian trees skin and produce spores at their tips had

Fungi, Ziser, Lecture Notes, 2011.3 61 Fungi, Ziser, Lecture Notes, 2011.3 62

when mature the sticky spores are are shot off

those that land on other flies continue the cycle

12. Future uses of fungi and fungal products

a. Fungi as & pesticides

eg. some have suggested using the fungus to be used as a biological control agent against drug crops in foreign countries

but: Fusarium is highly prone to ! may infect plants not intended

its use against marijuana crops in US has already been rejected because of this

eg. some fungi that parasitize arthropods are being investigated as potential “”

more than 50 fungal pathogens of insects are known

many are endemic and produce high mortality rates

not much luck yet, too many variables b. yeasts as food

a rich and easy to grow food source

high in B vitamins and proteins

1 acre ! 70 lbs of meat or milk protein (cow) ! 800 lbs of yeast protein

Fungi, Ziser, Lecture Notes, 2011.3 63