ENVU4E4 Alternative Exam Paper 2 – 3113228

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Question B:
Discuss different the role of symbiotic organisms in the supply of nitrogen (N) to plants.

Plan:
Mycorrhizal fungi – what are they and what do they do
- https://canvas.stir.ac.uk/courses/14614/pages/lecture-12-the-temperate-biome
Guest lecture – Paul Thomas
Entangled life
Mycorrhizas in ecosystem
Mycorrhizal symbiosis
Biogeochemistry (?) (pp 190-192)
Legume symbiosis (use article, https://www.jstor.org/stable/42947859 ) ), have got it
printed but not sure where it is.
- Leguminosae species form nodules but not all do
BNF Symbiosis
- Bacteria which have symbiotic relationship with plants to fix nitrogen in the nitrogen
cycle.

Sources:
See reading list for Paul’s references
https://librarysearch.stir.ac.uk/discovery/search?query=any,contains,mycorrhizal%20fungi&
cvtab=Everything&search_scope=MyInst_and_CI&vid=44UST_INST:VU1&offset=0
https://librarysearch.stir.ac.uk/discovery/search?query=any,contains,plant%20symbiosis&t
ab=LibraryCatalog&search_scope=ALMA_NO_JOURNALS&vid=44UST_INST:VU1&offset=0
Potential Sources:

AM AND EM
- Smith and Read, 2008 https://ebookcentral.proquest.com/lib/stir/detail.action?pq-
origsite=primo&docID=349538 (Book). 3rd edition, 1st out on loan (me).
 Essay:
Plants are unable to use atmospheric nitrogen, so the majority of plant species form
symbiotic relationships with many groups of organisms to obtain nutrients such as nitrogen
from the soil. It is mostly fungi and bacteria that plants have a symbiotic relationship with.
Fungi from the phylum Glomeromycota, Basidiomycota, and Ascomycota can all form
symbiotic mycorrhizal associations with plant roots, while bacteria from the genera
Bradyrhizobium, Rhizobium, and Frankia are species of nitrogen-fixing bacteria. Biological
nitrogen fixation is the conversion of raw atmospheric nitrogen (N 2) to ammonia (NH3),
which plants can use. Dark septate endophytes from the phyla Ascomycota also provide
nitrogen to the plant for nutritional purposes. (Gilbert et al, 2019).
Mycorrhizal, literally means fungus root, symbiosis is mutualistic, this is because both the
plant and the fungi benefit from each other. Plants receive nutrients and in exchange, the
fungi will receive carbohydrates from the plant as they can’t synthesize their own. There are
two main types of mycorrhizal association, arbuscular mycorrhizal (AM) and ectomycorrhizal
(EM). Mycorrhizal associations often connect many plants together under the soil due to the
extensive fungal networks they produce, a phenomenon known as the Wood Wide Web.
EXPAND -
https://www.jstor.org/stable/25609635?sid=primo&saml_data=eyJzYW1sVG9rZW4iOiI2OD
EzMzEwMS1iYzAyLTQ5MmYtOTI3OC01MTU4YWFmMDhhMTUiLCJpbnN0aXR1dGlvbklkcyI6
WyIzMWZjNzBmZC02N2IzLTQ5ZTUtYTI3YS1mNzBiN2RhYTNkMjkiXX0
In arbuscular mycorrhizal associations (AM) (also called Vesicular-arbuscular mycorrhiza
(Harley and Smith, 1983)) the fungi form intracellularly and penetrate the root cells forming
arbuscules with the root's cortical cells, which carry out nutrient exchange, within the root.
These mycorrhizal associations are found in 90% of all plant families and the colonization of
land relied on this relationship. Arbuscular mycorrhizas are found exclusively within the
phylum Glomeromycota (Schlesinger and Bernhardt, 2020). Some plants are capable of
forming both arbuscular and ectomycorrhizal associations. This dual relationship enhances
the nitrogen transfer that takes place (Harley and Smith, 1983).
Ectomycorrhizal fungi (EM), on the other hand, form extracellularly and form a coating
around plant roots and extend their Hartig nets made of hyphae throughout the soil
(Schlesinger and Bernhardt, 2020). They can only form in a limited number of species of
trees and shrubs and fungal species. Ectomycorrhizal associations have the added
advantage of being able to access insoluble organic forms of nitrogen which are inaccessible
to other types of mycorrhizal association such as arbuscular mycorrhizal and dark septate
root endopytes (Lambers et al, 2019).
Dark septate root endophytes (DSEs) are fungi in the phylum Ascomycota and order
Helotiales. They are commonly found in areas that have polar or alpine climates. Dark
septate root endophytes also form mutualistic relationships with plants, similar to those of
mycorrhizal fungi. It has been observed that dark septate endophytes can increase the
concentration of nitrogen by up to 103% compared to plants that do not form this
mutualistic relationship. This is due to the fact that the fungi are capable of transferring raw
nitrogen in the soils to the plants as part of the symbiotic relationship. It is also found the
biomass of these plants, specifically in the roots and shoots, is increased as nitrogen is a
critical nutrient in plant growth and development (Newsham, 2011).
Biological nitrogen fixation is the conversion of raw atmospheric nitrogen (N 2) to ammonia
(NH3), which plants can use. This process is carried out by many bacteria species which form
a symbiotic relationship with plant roots. There are four prerequisites that the relationship
has to meet to be successful in biological nitrogen fixation. These are: the bacteria has to be
able to form a symbiotic relationship with the plant, the plant must be able to provide
nutrients to the bacteria, they must be able to utilize oxygen as nitrogen fixation is an
aerobic process and the bacteria must be able to transport any nitrogen they fix to the plant
via its roots. The bacterial groups that can carry out nitrogen fixation are Bradyrhizobium,
Rhizobium, and Frankia. This symbiotic relationship is often found in species of Legumes
(Family: Leguminosae) and Actinorhizal (a group of plants within the angiosperms that are
characterised by the ability to form a symbiosis with Frankia bacteria) (Vessey et al, 2005).
This biological nitrogen fixation symbiosis has been observed between members of the
Leguminosae family and Rhizobium bacteria. In this symbiosis, the bacteria will form
nodules around the roots of the Leguminosae plant. These nodules will contain the bacteria
which will then be able to fix the nitrogen and convert it into ammonia (Bryan et al, 1996).
These nodules also contain the enzyme nitrogenase which facilitates the biological nitrogen
fixation reaction. The legumes will also limit the growth and reproductive capacity of the
bacteria, if the bacterial levels get too high, the bacteria will use the ammonia they produce
rather than transporting it to the plant. In return, the legumes will provide essential
nutrients to the Rhizobium, making this a mutualistic relationship (Downie, 2014).

Actinorhizal plants can form symbiotic relationships with gram positive bacteria of the genus
Frankia. This is the only genus of bacteria that this group of angiosperms can form a
relationship with. Like the legumes described above, Frankia also form bacteria containing
nodules for the purposes of biological nitrogen fixation (Froussat et al, 2016).

References:
Gibert, A., Tozer, W. and Westoby, M. (2019), Plant performance response to eight different
types of symbiosis. New Phytol, 222: 526-542. Available at:
https://doi.org/10.1111/nph.15392
Schlesinger, W. H. & Bernhardt, E. S. (2020) Biogeochemistry: an analysis of global change.
Fourth edition. London; Academic Press. Available at: https://www-sciencedirect-
com.ezproxy-s1.stir.ac.uk/book/9780128146088/biogeochemistry

Newsham, K.K. (2011), A meta-analysis of plant responses to dark septate root endophytes.
New Phytologist, 190: 783-793. Available at: https://doi.org/10.1111/j.1469-
8137.2010.03611.x
Vessey, J.K., Pawlowski, K. & Bergman, B. (2005) Root-based N2-fixing Symbioses: Legumes,
Actinorhizal Plants, Parasponia sp. and Cycads. Plant Soil 274, 51–78. Available at:
https://doi.org/10.1007/s11104-005-5881-5
Bryan, J.A et al (1996) Toward a new concept of the evolution of symbiotic nitrogen fixation
in the Leguminosae, Plant and Soil 186 pp151-159 Available at:
https://www.jstor.org/stable/42947859?seq=7
Froussart, E., Bonneau, J., Franche, C. et al. (2016) Recent advances in actinorhizal symbiosis
signaling. Plant Mol Biol 90, 613–622. Available at: https://doi.org/10.1007/s11103-016-
0450-2
Downie, J.A, (2014). Legume Nodulation, Current Biology 24(5) pp R184-R190 Available at:
https://www.sciencedirect.com/science/article/pii/S096098221400061X?pes=vor
Harley, J. L. & Smith, S. E. (1983) Mycorrhizal symbiosis 1st edition. London: Academic Press.
Lambers, H. et al (2008). Plant nutrient-acquisition strategies change with soil age, Trends in
Ecology and Evolution 23(2) pp 95-103 Available at:
https://www.sciencedirect.com/science/article/pii/S0169534707003576