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Actinorhizal plant
From Wikipedia, the free encyclopedia

Actinorhizal plants are a group of angiosperms characterized by their ability to

form a symbiosis with the nitrogen xing actinobacteria Frankia. This association
leads to the formation of nitrogen-xing root nodules.

Contents
1 Classication
2 Distribution and ecology
3 Evolutionary origin
4 The symbiotic nodules
5 Notes
6 References
7 External links

Classication
Actinorhizal plants are dicotyledons distributed among three angiosperm orders, 8
families and 24 genera:[1]

Cucurbitales Fagales Rosales

Coriariaceae Betulaceae Elaeagnaceae
Coriaria Alnus Elaeagnus
Datiscaceae Casuarinaceae Hippophae
Datisca Allocasuarina Shepherdia
Casuarina Rhamnaceae
Ceuthostoma Colletia
Gymnostoma Discaria
Myricaceae Kentrothamnus
Comptonia Retanilla
Myrica Talguenea
(including Trevoa
Morella) Ceanothus
Rosaceae
Cercocarpus
Chamaebatia
Cowania
Dryas
Purshia

These three orders form a single clade within the Rosids, which is a sister taxon to
the other major nitrogen-xing order, the Fabales. All actinorhizal species are trees

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or shrubs, except for the genus Datisca.

Many are common plants in temperate regions like alder, bayberry, sweetfern,
Avens, mountain misery and Coriaria. Some Elaeagnus species and Sea-buckthorns
produce edible fruit. In tropical regions, Casuarinas are widely cultivated.

Distribution and ecology

Actinorhizal plants are found on all continents
except for Antarctica. Their ability to form nitrogen-
xing nodules confers a selective advantage in poor
soils. Most actinorhizal plants are therefore pioneer
species that colonize young soils where available
nitrogen is scarce like moraines, volcanic ows or
sand dunes.[2] Being among the rst species to
The distribution of
colonize these disturbed environments, actinorhizal
shrubs and trees play a critical role, enriching the actinorhizal plants.
soil and enabling the establishment of other species
in an ecological succession.[1][2] Actinorhizal plants like alders are also common in
the riparian forest.[2]

Actinorhizal plants are the major contributors to nitrogen xation in broad areas of
the world, and are particularly important in temperate forests.[1] The nitrogen
xation rate measured for some alder species is as high as 300kg of N2/ha/year,
close to the highest rate reported in legumes.[3]

Evolutionary origin
No fossil records are available concerning nodules, but fossil pollen of plants
similar to modern actinorhizal species has been found in sediments deposited 87
million years ago. The origin of the symbiotic association remains uncertain. The
ability to associate with Frankia is a polyphyletic character and has probably
evolved independently in dierent clades.[4] Nevertheless, actinorhizal plants and
Legumes, the two major nitrogen-xing groups of plants share a relatively close
ancestor, as they all part of a clade within the rosids which is often called the
nitrogen-xing clade. This ancestor may have developed a "predisposition" to enter
in symbiosis with nitrogen xing bacteria and this led to the independent
acquisition of symbiotic abilities by ancestors of the actinorhizal and Legume
species. The genetic program used to establish the symbiosis has probably
recruited elements of the arbuscular mycorrhizal symbioses, a much older and
widely distributed symbiotic association between plants and fungi. [5]

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Evolutionary
origin of nitrogen-
xing nodulation

The symbiotic nodules

As in legumes, nodulation is favored by nitrogen deprivation and is inhibited by
high nitrogen concentrations. Depending on the plant species, two mechanisms of
infection have been described: The rst is observed in casuarinas or alders and is
called root hair infection. In this case the infection begins with an intracellular
infection of a root hair and is followed by the formation of a primitive symbiotic
organ lacking any particular organization, a prenodule.[6] The second mechanism
of infection is called intercellular entry and is well described in Discaria species. In
this case bacteria penetrate the root extracellularly, growing between epidermal
cells then between cortical cells. Later on Frankia becomes intracellular but no
prenodule is formed. In both cases the infection leads to cell divisions in the
pericycle and the formation of a new organ consisting of several lobes anatomically
similar to a lateral root. This organ is the actinorhizal nodule also called
actinorhizae. Cortical cells of the nodule are invaded by Frankia laments coming
from the site of infection or the prenodule. Actinorhizal nodules have generally an
indeterminate growth, new cells are therefore continually produced at the apex
and successively become infected. Mature cells of the nodule are lled with
bacterial laments that actively x nitrogen.

Little information is available concerning the mechanisms leading to nodulation.

No equivalent of the rhizobial Nod factors have been found, but several genes
known to participate in the formation and functioning of Legume nodules (coding
for heamoglobin and other nodulins) are also found in actinorhizal plants where
they are supposed to play similar roles.[7] The lack of genetic tools in Frankia and
in actinorhizal species was the main factor explaining such a poor understating of
this symbiosis, but the recent sequencing of 3 Frankia genomes [8] and the
development of RNAi and genomic tools in actinorhizal species [9][10] should help
to a far better understanding in the following years.

Notes
1. Wall 2000 3. Zavitovski & Newton 5. Kistner & Parniske
2. Schwintzer & Tjepkema 1968 2002
1990 4. Benson & Clawson 6. Laplaze 2000
2000 7. Vessey 2003

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8. Normand 2006 9. Hocher 2006 10. Gherbi 2008

References
Wall, L. (2000), "The actinorhizal symbiosis", Journal of Plant Growth and
Regulation, 19 (2): 167182, doi:10.1007/s003440000027, PMID 11038226

Schwintzer, C. R.; Tjepkema, J. (1990), The Biology of Frankia and

Actinorhizal Plants, Academic Press, ISBN 012633210X

Benson, D. R.; Clawson, M. L. (2000), "Evolution of the actinorhizal plant

nitrogen-xing symbiosis", in Triplett, E., Prokaryotic Nitrogen Fixation: A
Model System for the Analysis of a Biological Process, Norfolk, UK: Horizon
Scientic Press, pp.207224, ISBN 1-898486-19-0

Zavitovski, J.; Newton, M. (1968), "Ecological importance of snowbrush

Ceanothus velutinus in the Oregon cascade", Ecology, 49 (6): 11341145,
doi:10.2307/1934497, JSTOR 1934497

Kistner, C.; Parniske, M. (2002), "Evolution of signal transduction in

intracellular symbiosis", Trends in Plant Science, 7 (11): 511518,
doi:10.1016/S1360-1385(02)02356-7, PMID 12417152

Laplaze, L.; Duhoux, E.; Franche, C.; Frutz, T.; Svistoono, S.; Bisseling, T.;
Bogusz, D.; Pawlowski, K. (2000), "Casuarina glauca prenodule cells display
the same dierentiation as the corresponding nodule cells" (PDF), Molecular
Plant-Microbe Interactions, 13: 107112, doi:10.1094/MPMI.2000.13.1.107,
PMID 10656591

Vessey, J. K.; Pawlowski, K.; Bergman, B. (2005), "Root-based N 2-xing

symbioses: Legumes, actinorhizal plants, Parasponia sp. and cycads", Plant
and soil, 266 (1): 205230, doi:10.1007/s11104-005-0871-1

Gherbi, H.; Markmann, K.; Svistoono, S.; Estevan, J.; Autran, D.; Giczey, G.;
Auguy, F.; Pret, B.; Laplaze, L.; Franche, C.; Parniske, M.; Bogusz, D. (2008),
"SymRK denes a common genetic basis for plant root endosymbioses with
arbuscular mycorrhiza fungi, rhizobia, and Frankia bacteria", Proceedings of
the National Academy of Sciences, 105 (12): 49284932,
doi:10.1073/pnas.0710618105, PMC 2290763 , PMID 18316735

Hocher, V.; Auguy, F.; Argout, X.; Laplaze, L.; Franche, C.; Bogusz, D. (2006),
"Expressed sequence-tag analysis in Casuarina glauca actinorhizal nodule and
root", New Phytologist, 169 (4): 681688,
doi:10.1111/j.1469-8137.2006.01644.x, PMID 16441749

Normand, P.; Lapierre, P.; Tisa, L. S.; Gogarten, J. P.; Alloisio, N.; Bagnarol, E.;
Bassi, C. A.; Berry, A. M.; Bickhart, D. M.; Choisne, N.; Couloux, A.;
Cournoyer, B.; Cruveiller, S.; Daubin, V.; Demange, N.; Francino, M. P.;
Goltsman, E.; Huang, Y.; Kopp, O. R.; Labarre, L.; Lapidus, A.; Lavire, C.;
Marechal, J.; Martinez, M.; Mastronunzio, J. E.; Mullin, B. C.; Niemann, J.;

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Pujic, P.; Rawnsley, T.; Rouy, Z. (2006). "Genome characteristics of

facultatively symbiotic Frankia sp. Strains reect host range and host plant
biogeography". Genome Research. 17 (1): 715. doi:10.1101/gr.5798407.
PMC 1716269 . PMID 17151343.

External links
Frankia and Actinorhizal plant Website
Wikimedia Commons
(http://web.uconn.edu/mcbsta/benson/Frankia has media related to
/FrankiaHome.htm) Actinorhizal plant.

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Categories: Biogeochemical cycle Nitrogen Nitrogen cycle Soil biology

Symbiosis

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