
|
White Pine seedlings grown in sterile conditions (left),
White Pine seedlings grown in forest soil with mychorrhizae.
Photo courtesy of www.msu.educourseisb202 |
About 90% of extant land plants form symbioses with fungi (Cairney 2000). In phototrophs that have secondarily reverted back to the aquatic environment, their ability to form relationships with fungi is lost, providing more evidence that phototrophs evolved onto land with the aid of fungal associations (Pirozynski and Malloch 1988). Today, Atlantic white cedar fluctuate their levels of mycorrhizal "infection" during drought and flooding (Cantelmo and Ehrenfeld 1999), which indicate how plants regulate their fungal needs and also may be reminiscent of the first plant’s journey to land. Also, plants that do not form mycorrhizal relations are mostly in highly disturbed habitats or in wet or aquatic habitats where mineral resources are adequate and the diffusion of oxygen prohibits the growth of fungi (Fitter and Peat 1993). Even algae living in tidal zones form mycophycobiosis during low tides to cope with the desiccating stress (Kohlmeyer and Kohlmeyer 1979). Lastly, fossil evidence agrees with molecular data in that arbuscular mycorrhizae form a monopyletic group and arose around 353-462 million years ago, around the same time plants colonized land (Redecker et al 2000).
|
Arbuscular mycorrhizae.
Photo courtesy of terroirists.nettagmicrobiology
|
Types of Mycorrhizae
Arbuscular mycorrhizae (AM) are mychorrhizae that actually penetrate the plant's cortical tissue and are the most numerous type of mycorrhizae in present-day plants. However, AM consists of only 130 species (Morton 1990) and shows generality to plant hosts (Perry 1998). AM’s include fungi in the phylum Zygomycota, more specifically the order Glomales, and the plant hosts include most angiosperms, some gymnosperms, Pteridophytes and various lower plants (Smith and Read 1997). These AM associations occur mostly at mid-latitudes where phosphorus for plant consumption is limited (Read 1991). Fossil and molecular evidence significantly indicate that the ancestors of extant plants formed AM relationships (Cairn 2000) and today’s plants who do not have AM have lost their relationship with their ancestral host plant (Barker et al. 1998). Gehrig et al. (1996) have found a fungus endocytobiont, Geosiphon pyriforme, to be related to an ancestral form of Glomales, thus forming a monophyletic group, and is probably most like the Glomus fungi that first aided plants to adapt to land life. In addition, molecular clock analysis suggests that the phylogenetic radiation of ancient Glomales paralleled the colonization of land (Redecker et al 2000).
 |
Ectomycorrhizae.
Photo courtesy of farm3.static.flickr.com |
Another type of mychorrhizae, Ectomycorrhizae (ECM), form a sheath around the outside of the root of the plant and are only on woody trees and shrubs (Cairney 2000). ECM fungi permit their host to acquire phosphorus, nitrogen and organic material (Read 1991). ECM have a great importance in shaping the ecosystems of forests and involve fungi that belong to the phylums Basidiomycota, Ascomycota and Zygomycota (Cairney 2000). Interestingly, however, ECM extant plants can also form AM associations, depending on the soil conditions (Smith and Read 1997). In addition, some ECM capable plants only form AM associations at the seedling stage, providing even more evidence that all land plants that exist at the present evolved from an ancestral AM condition (Cairney 2000).
Ericoid mycorrhizae (ERM) occur in extremely nutrient poor soils and at high latitudes and altitudes (Read 1991). The fungi in this association have extensive coils of hyphae that cover the epidermal cells of the plant host and involve specifically Ericad plants (the Heather family) and Ascomycete fungi (Cairney 2000). ERM’s have good saprotrophic abilities permitting them to provide nitrogen and phosphorus and tolerate toxic cations that are in acidic soils (Smith and Read 1997).
 |
Photo courtesy of the botany department at WVU |
Conclusion
Coevolution is the reciprocal genetic change among a species or populations (Thompson 1999), while parallel evolution is the result of similar pressures acting on species and results in similar yet independent outcomes. Which evolutionary process acted on plants and fungi in mychorrizal associations? Currently evidence for pure coevolution is lacking (Cairney 2000). However, Juenger and Bergelson (1998) propose that the coevolution is more “diffuse” and at a “guild” level of selection. Cairney (2000) has looked at this very question in his paper, The Evolution of Mycorrhiza Systems. He believes that coevolution did occur, but he sees no evidence that gene-for-gene coevolution is occurring in extant species and that it is simply parallel evolution at the present day.
Despite the lack of direct evidence that coevolution is occurring today in mycorrhizae, I think that coevolution is indeed occurring today. If coevolution were a continuum, then I would suggest the exant fungal and plant species are to a lesser degree coevolving because I believe there is some gene-for-gene reciprocation. Mycorrhizal associations can help a plant survive and reproduce, excluding certain genomes that cannot fully exploit this symbiosis. Generalities occur in fungal associations because of common ancestral lineages and does not infer that parallel evolution is occurring. It is exhibited that in certain environmental conditions certain mycorrhizal associations take place, which could support parallel evolution. Perhaps it is not, since a basis for the two organisms to come together in symbiosis must have been established at some point to permit the relationship. I suggest that mycorrhizal associations are still coevolving, to a degree, in which genetic give and take must take place. It is important when creating fungal phylogenies of those that form mycorrhizae to include plant hosts and vice versa, for mycorrhizae associations were and are the essential associations for plants to survive.
With the current prevalence and severity of habitat destruction and the overpopulation that permits limited resources, soil structure will never be perfect everywhere on Earth. However, for native plants and ecosystems to become or remain healthy and supporting, mycorrhizae are and will be the mechanism of survival for all life forms. In the future, man will foresee and artificially create these associations for habitat restoration and crop production and this application will be an important factor that will enable humans to feed one another and rebuild plundered ecosystems.
Barker, S.J. Tagu, D. Delp, G. 1998. Regulation of root and fungal morphogenesis in mycorrhizal symbioses. Plant Physiol. 116: 1201-1207
Cairney, J.W.G. 2000. “Evolution of mycorrhiza systems.” Natuwissenschaaften. 87:467-475
Cantelmo, A.J. and Ehrenfeld, J.G. 1999. Effects of microtopography on mycorrhizal infection in Atlantic white cedar pine. Mycorrhiza. 8: 175-180
Fitter, A.H. and Peat, H.J. The distribution of arbuscular mycorrhizas in the British flora. New Phytol. 125, 845-854
Fitter, A.H. and Moyerson, B. 1996. Evolutionary trends in root-microbe symbioses. Phil Trans R Soc Lond B. 351:1367-1375
Gehrig, H. Schubler, A. Kluge, M. 1996. Geosiphon pyriforme, a fungus forming endocytobiosis with Nostoc, is an ancestral member of Glomales: evidence by SSU rRNA analysis. J Mol Evol. 43: 71-81
Hawksworth, D.I. 1988. Coevolution of fungi with algae and cyanobacteria in lichen symbioses. Coevolution of Fungi with Plants and Animals. Academic Press. 125-148
Heijden, M.G.A. van der. Boller T. Wiemken A. Sanders I.R. 1998. Different arbuscular mycorrhizal fungal species are potential determinants of plant community structure. Ecology. 79: 2082-2091.
Juenger, T. Bergelson, J.1998. Pairwise versus diffuse natural selection and the multiple herbivores of scarlet gilia, Ipomopsis aggregata. Evolution. 52: 1583-1592
Kohlmeyer, E. and Kohlmeyer J. 1979. Marine Mycology: the Higher Fungi, Academic Press
Morton, J.B. Benny, G.L. 1990. Revised classification of arbuscular mycorrhizal fungi. Mycotaxon. 37:471-492
Perry, David. 1998. A movable feast: the evolution of resource sharing in plant-fungus communities.Trends in Ecology and Evolution. Vol.13, issue 11: 432-434
Pirozynski, KA and Malloch, DW. 1975. The origin of land plants: a matter of mycotrophism. BioSystems. 6:153-164
Read, D.J. 1991. Mycorrhizas in ecosystems. Experientia. 47:376-309
Thompson, J.N. 1999 The evolution of species interactions. Science. 284:2116-2118
Redecker, D. Kodner, R. Graham, L.E. 2000. Glomalean fungi from the Ordovician. Science. 289: 1920-1921.
Simon et al. 1993. Origin and diversification of endomycorrhizal fungi and the coincidence with vascular plants. Nature 363: 67-69
Smith M.D. and Read D.J. 1997. Mycorrhizal symbiosis. Academic Press, London
Thompson, J.N. 1999. The evolution of species interactions. Science. 284: 2116-2118.