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= Flower-Mimicking Rust Fungi =
= Flower-Mimicking Rust Fungi =
Several fungal plant pathogens infect a host plant and alter its morphology in some fashion, but few alter the flowers of their hosts. Flower-mimicking rust fungi infect their host which results in the creation of pseudoflowers made by the host to aid the fungi in gamete dispersal for reproduction. These rust fungi are parasites, requiring a living host plant i which to carry out their 5 life stages.
Several fungal plant pathogens infect a host [[plant]] and alter its morphology in some fashion, but few alter the [[Flower|flowers]] of their hosts. Flower-mimicking [[Rust (fungus)|rust fungi]] infect their host which results in the creation of pseudoflowers made by the host plant to aid the fungi in gamete dispersal for sexual reproduction. These rust fungi are parasites, requiring a living host plant in which to carry out their five life stages.


== Species ==
== Species ==
Two species of rust fungi are known to infect and alter their plant host's flower morphology, resulting in the formation of structures on the plant that look similar to true flowers. These two species are ''[[Puccinia monoica]]'' and ''[[Uromyces pisi-sativi|Uromyces pisi]]''. Both species are in the same order of fungus, Pucciniales. <ref>http://website.nbm-mnb.ca/mycologywebpages/NaturalHistoryOfFungi/Pucciniales.html</ref>
Two species of rust fungi are known to infect and alter their plant host's flower morphology, resulting in the formation of structures on the plant that look similar to true flowers.<ref name=":3" /> These two species are ''[[Puccinia monoica]]'' and ''[[Uromyces pisi-sativi|Uromyces pisi]]''. Both species are in the same order of fungi, Pucciniales.<ref name=":3">http://website.nbm-mnb.ca/mycologywebpages/NaturalHistoryOfFungi/Pucciniales.html</ref>


=== Puccinia monoica ===
=== Puccinia monoica ===
''Puccinia monoica'' is a crown rust that infects a range of species in the botanical genus ''[[Arabis]]'' <nowiki/>, within the family Brassicaceae<ref name="Roy1993">Roy, B. A. "Floral Mimicry by a Plant Pathogen." Nature 362 (1993): 56 - 58.</ref> . It also infects species outside of ''Arabis'', notably ''[[Boechera stricta]]''.
''Puccia monoica'' is a crown rust that infects a range of species in the botanical genus ''[[Arabis]],'' <nowiki/>within the family Brassicaceae, affecting 11 Brassicaceae species overall.<ref name="Roy1993">Roy, B. A. "Floral Mimicry by a Plant Pathogen." Nature 362 (1993): 56 - 58.</ref> It also infects species outside of ''Arabis'', notably ''[[Boechera stricta]]'', which also is in the family Brassicaceae.


=== Uromyces pisi ===
=== Uromyces pisi ===
''Uromyces pisi'' infects a perennial ornamental plant, ''[[Euphorbia cyparissias]]''<nowiki/><ref name=":0">Pfunder, Monika, and Bitty A. Roy. "Pollinator-mediated interactions between a pathogenic fungus, Uromyces pisi (Pucciniaceae), and its host plant, Euphorbia cyparissias (Euphorbiaceae)." American Journal of Botany 87 (2000): 48 - 55.</ref> . It is also known to infect plants in the family Fabaceae<ref>{{Cite journal|last=Pfunder|first=Monika|last2=Schürch|first2=Stéphanie|last3=Roy|first3=Barbara A.|title=Sequence variation and geographic distribution of pseudoflower-forming rust fungi (Uromyces pisi s. lat.) on Euphorbia cyparissias|url=https://doi.org/10.1017/S0953756200003208|journal=Mycological Research|volume=105|issue=1|pages=57–66|doi=10.1017/s0953756200003208}}</ref>.
''Uromyces pisi'' infects a perennial ornamental plant, ''[[Euphorbia cyparissias]]''<nowiki/><ref name=":0">Pfunder, Monika, and Bitty A. Roy. "Pollinator-mediated interactions between a pathogenic fungus, Uromyces pisi (Pucciniaceae), and its host plant, Euphorbia cyparissias (Euphorbiaceae)." American Journal of Botany 87 (2000): 48 - 55.</ref>, within the family [[Euphorbiaceae]]. It is also known to infect plants in the family Fabaceae.<ref>{{Cite journal|last=Pfunder|first=Monika|last2=Schürch|first2=Stéphanie|last3=Roy|first3=Barbara A.|title=Sequence variation and geographic distribution of pseudoflower-forming rust fungi (Uromyces pisi s. lat.) on Euphorbia cyparissias|url=https://doi.org/10.1017/S0953756200003208|journal=Mycological Research|volume=105|issue=1|pages=57–66|doi=10.1017/s0953756200003208}}</ref>


== Characteristics ==
== Characteristics ==


=== Pseudoflowers ===
=== Pseudoflowers ===
Pathogenic rust fungi infect their host plant and induce the formation of pseudoflowers. These pseudoflowers are modified leaves that resemble flowers of another unrelated plant species.<ref name="Roy1993" /> The pseudoflowers differ from the host plant flowers in shape, size, color, scent, and nectar produced. These pseudoflowers are formed as yellow rosettes and produce a sweet-smelling nectar. The mimicry expressed here is similar to [[Dodsonian mimicry]] in which reproductive flowers are mimicked by another species. However, in Dodsonian mimicry, nectar is not produced by the impostor flower. The pseudoflowers created by these rust fungi create a sweet nectar that attract insect pollinators.
Pathogenic rust fungi infect their host plant and induce the formation of pseudoflowers. These pseudoflowers are modified leaves that resemble flowers of another unrelated plant species.<ref name="Roy1993" /> The pseudoflowers differ from the host plant flowers in shape, size, color, scent, and nectar produced. These pseudoflowers are formed as yellow rosettes with embedded fungal spermagonia and produce a sweet-smelling nectar to attract and reward insect pollinators.<ref name=":1" /> The mimicry expressed here is similar to [[Dodsonian mimicry]] in which reproductive flowers are mimicked by another species. However, in Dodsonian mimicry, nectar is not produced by the impostor flower.


=== Scent Chemistry ===
=== Scent Chemistry ===
The fragrance of the pseudoflowers produced by rust fungi is one of the main characteristics that attracts insect pollinators. The chemical compounds that form the fragrance of the fungal pseudoflowers differs between species. Mass spectrometry and gas chromatography analyses have allowed identification of different fragrances of these pseudoflowers. Curiously, some fragrances are found to be chemically similar to non-host flowers rather than to host flowers and host vegetation. Fungal pseudoflowers are mainly composed of aromatic alcohols, aldehydes and esters<ref>Raguso, Robert A., and Barbara A. Roy. "‘Floral’scent production by Puccinia rust fungi that mimic flowers." Molecular Ecology 7.9 (1998): 1127-1136.</ref>. In P. monoica, the fungal pseudoflower scent is made of aromatic compounds including phenylacetaldehyde and 2-phenylethyl alcohol. In Uromyces pisi, the fungal pseudoflower scent is made of compounds including indole and jasmine lactone.<ref>{{Cite journal|last=Kaiser|first=Roman|date=2006-02-10|title=Flowers and Fungi Use Scents to Mimic Each Other|url=http://science.sciencemag.org/content/311/5762/806|journal=Science|language=en|volume=311|issue=5762|pages=806–807|doi=10.1126/science.1119499|issn=0036-8075|pmid=16469916}}</ref>
The fragrance of the pseudoflowers produced by rust fungi is one of the main characteristics that attracts insect pollinators.<ref name=":1" /> The chemical compounds that form the fragrance of the fungal pseudoflowers differs between species. Mass spectrometry and gas chromatography analyses have allowed identification of different fragrances of these pseudoflowers.<ref name=":1" /> Curiously, some fragrances are found to be chemically similar to non-host flowers rather than to host flowers and host vegetation. ''P. monoica'' pseudoflowers are mainly composed of aromatic alcohols, aldehydes and esters.<ref name=":1">Raguso, Robert A., and Barbara A. Roy. "‘Floral’scent production by Puccinia rust fungi that mimic flowers." Molecular Ecology 7.9 (1998): 1127-1136.</ref> In ''P. monoica'', the fungal pseudoflower scent is dominated by aromatic compounds including phenylacetaldehyde and 2-phenylethyl alcohol. Comparatively, the scent of flowers of ''Arabis'' hosts is dominated by fatty-acid derived products. ''P. monoica's'' pseudoflowers are thought to be most similar to unspecialized [[Leontopodium nivale|Edelwiess]] flowers due to similarities in scent compounds.
In Uromyces pisi, the fungal pseudoflower scent is made of compounds including indole and jasmine lactone.<ref name=":2">{{Cite journal|last=Kaiser|first=Roman|date=2006-02-10|title=Flowers and Fungi Use Scents to Mimic Each Other|url=http://science.sciencemag.org/content/311/5762/806|journal=Science|language=en|volume=311|issue=5762|pages=806–807|doi=10.1126/science.1119499|issn=0036-8075|pmid=16469916}}</ref> The scent of ''E. cyparissias'' pseudoflowers is known to vary based on geographic location, ranging from a floral-fruity smell to a spermy scent.<ref name=":2" />

== Life Cycle ==
== Life Cycle ==


=== Puccinia monoica life cycle ===
=== Puccinia monoica life cycle ===
Puccinia monoica has a macrocyclic, heteroecious life cycle. Macrocyclic refers to the fungus succesively completing 5 spore states in its life. Heteroecious refers to having two complementary mating types housed in separate organisms. The reproduction of a heteroecious species relies upon a dispersal mechanism in order for outcrossing between the mating types to occur. The life cycle of Puccinia monoica is described in more detail [[Puccinia monoica|here]].
''P. monoica'' has a macrocyclic, heteroecious life cycle. Macrocyclic refers to the fungus succesively completing 5 spore states in its life. Heteroecious refers to having two complementary mating types housed in separate organisms. The reproduction of a heteroecious species relies upon a dispersal mechanism in order for outcrossing between the mating types to occur. The life cycle of ''P. monoica'' is described in more detail [[Puccinia monoica|here]].


=== Uromyces pisi life cycle ===
=== Uromyces pisi life cycle ===
Uromyces pisi also has a macrocyclic, heteroecious life cycle<ref>Barilli E., Sillero J.C., Prats E., Rubiales D. "Resistance to rusts (Uromyces pisi and U. viciae-fabae) in pea." Czech J. Genet. Plant Breed 50 (2014): 135–143.</ref>, meaning it successively completes 5 spore stages in a complete life cycle. Like P. monoica, outcrossing between mating types depends on a dispersal mechanism.
''U. pisi'' also has a macrocyclic, heteroecious life cycle.<ref>Barilli E., Sillero J.C., Prats E., Rubiales D. "Resistance to rusts (Uromyces pisi and U. viciae-fabae) in pea." Czech J. Genet. Plant Breed 50 (2014): 135–143.</ref> meaning it successively completes five spore stages in a complete life cycle. The five spore stages created are: [[Pycniospore|Pycniospores]] (spermatia), [[Aeciospore|Aeciospores]], [[Urediniospore|Urediniospores]], [[Teliospore|Teliospores]], and [[Basidiospore|Basidiospores]]. Like ''P. monoica'', outcrossing between mating types depends on a dispersal mechanism by an insect pollinator. ''U. pisi'' alternates between two plant hosts to proceed through its life cycle. E. cyparissias hosts the fungal spermatia and aeciospores, while a species in the family [[Fabaceae]] is host to the other spore types.<ref>GAUMANN, E. Die Rostpilze Mitteleuropas. Beitrage zur Kryp-togamenflora der Schweiz Band 12 (1959): 1–1407.</ref>


== Ecology ==
== Ecology ==


=== Pseudoflower-insect interactions ===
=== Pseudoflower-insect interactions ===
Rust fungi reproduction depends on insect visitation, analogous to pollination in flowers. Insects transfer spermatia (gametes) between different mating types of infected plants. The presence of the pseudoflower in plant communities may have consequences other than enhance the reproduction of the fungus. Studies in ''P. monoica'' demonstrated that insect visitation occurs more often when plants present a mixture of both pseudoflowers and natural occurring flowers, rather than when they occur alone in the plant. Combination of real flowers and pseudoflowers in plants increase the floral display through and apparent higher density, it also produces complementary attraction cues and complementary pollinators rewards, thus it has been proposed that they play a role in the pollination dynamics between insect and plants.<ref>{{Cite journal|last=Roy|first=B. A.|date=1994-03-01|title=The Effects of Pathogen-Induced Pseudoflowers and Buttercups on Each Other's Insect Visitation|url=http://onlinelibrary.wiley.com/doi/10.2307/1939539/abstract|journal=Ecology|language=en|volume=75|issue=2|pages=352–358|doi=10.2307/1939539|issn=1939-9170}}</ref> In ''U. pisi'', insect pollinator visits are also necessary in order for two mating types of ''U. pisi'' to interact and result in fertilization.<ref name=":0" />
In U. pisi, insect pollinator visits are necessary in order for two mating types of U. pisi to interact and result in fertilization<ref name=":0" />.

Pseudoflowers are visited by [[Halictidae|halictid]], [[Andrenidae|andrenic]], [[Bumblebee|bumble]] and [[Oligolecty|oligolectic]] bees, muscid flies, and ants.<ref>Roy, B.A.Raguso, R.A.Olfactory versus visual cues in a floral mimicry system''Oecologia''109 1997414426</ref>


== See also ==
== See also ==
Line 41: Line 46:


== References ==
== References ==
- Roy, B. A. "The effects of pathogen‐induced pseudoflowers and buttercups on each other's insect visitation." Ecology 75.2 (1994): 352-358

- Pfunder, Monika, and Bitty A. Roy. "Fungal pseudoflowers can influence the fecundity of insect-pollinated flowers on Euphorbia cyparissias." Botanica Helvetica 116 (2006): 149 - 158.

- Ngugi, H. K., and Harald Scherm. "Mimicry in plant-parasitic fungi." FEMS Microbiology Letters 257 (2006): 171 - 176.

- Roy, B. A. "Floral Mimicry by a Plant Pathogen." Nature 362 (1993): 56 - 58.
Cano, L.M., et al. "Major Transcriptome Reprogramming Underlies Floral Mimicry Induced by the Rust Fungus Puccinia monoica in Boechera stricta." Plos ONE 8(9) (2013): 1 - 16.
<references />
<references />

Latest revision as of 06:14, 19 December 2017

Flower-Mimicking Rust Fungi

[edit]

Several fungal plant pathogens infect a host plant and alter its morphology in some fashion, but few alter the flowers of their hosts. Flower-mimicking rust fungi infect their host which results in the creation of pseudoflowers made by the host plant to aid the fungi in gamete dispersal for sexual reproduction. These rust fungi are parasites, requiring a living host plant in which to carry out their five life stages.

Species

[edit]

Two species of rust fungi are known to infect and alter their plant host's flower morphology, resulting in the formation of structures on the plant that look similar to true flowers.[1] These two species are Puccinia monoica and Uromyces pisi. Both species are in the same order of fungi, Pucciniales.[1]

Puccinia monoica

[edit]

Puccia monoica is a crown rust that infects a range of species in the botanical genus Arabis, within the family Brassicaceae, affecting 11 Brassicaceae species overall.[2] It also infects species outside of Arabis, notably Boechera stricta, which also is in the family Brassicaceae.

Uromyces pisi

[edit]

Uromyces pisi infects a perennial ornamental plant, Euphorbia cyparissias[3], within the family Euphorbiaceae. It is also known to infect plants in the family Fabaceae.[4]

Characteristics

[edit]

Pseudoflowers

[edit]

Pathogenic rust fungi infect their host plant and induce the formation of pseudoflowers. These pseudoflowers are modified leaves that resemble flowers of another unrelated plant species.[2] The pseudoflowers differ from the host plant flowers in shape, size, color, scent, and nectar produced. These pseudoflowers are formed as yellow rosettes with embedded fungal spermagonia and produce a sweet-smelling nectar to attract and reward insect pollinators.[5] The mimicry expressed here is similar to Dodsonian mimicry in which reproductive flowers are mimicked by another species. However, in Dodsonian mimicry, nectar is not produced by the impostor flower.

Scent Chemistry

[edit]

The fragrance of the pseudoflowers produced by rust fungi is one of the main characteristics that attracts insect pollinators.[5] The chemical compounds that form the fragrance of the fungal pseudoflowers differs between species. Mass spectrometry and gas chromatography analyses have allowed identification of different fragrances of these pseudoflowers.[5] Curiously, some fragrances are found to be chemically similar to non-host flowers rather than to host flowers and host vegetation. P. monoica pseudoflowers are mainly composed of aromatic alcohols, aldehydes and esters.[5] In P. monoica, the fungal pseudoflower scent is dominated by aromatic compounds including phenylacetaldehyde and 2-phenylethyl alcohol. Comparatively, the scent of flowers of Arabis hosts is dominated by fatty-acid derived products. P. monoica's pseudoflowers are thought to be most similar to unspecialized Edelwiess flowers due to similarities in scent compounds.

In Uromyces pisi, the fungal pseudoflower scent is made of compounds including indole and jasmine lactone.[6] The scent of E. cyparissias pseudoflowers is known to vary based on geographic location, ranging from a floral-fruity smell to a spermy scent.[6]

Life Cycle

[edit]

Puccinia monoica life cycle

[edit]

P. monoica has a macrocyclic, heteroecious life cycle. Macrocyclic refers to the fungus succesively completing 5 spore states in its life. Heteroecious refers to having two complementary mating types housed in separate organisms. The reproduction of a heteroecious species relies upon a dispersal mechanism in order for outcrossing between the mating types to occur. The life cycle of P. monoica is described in more detail here.

Uromyces pisi life cycle

[edit]

U. pisi also has a macrocyclic, heteroecious life cycle.[7] meaning it successively completes five spore stages in a complete life cycle. The five spore stages created are: Pycniospores (spermatia), Aeciospores, Urediniospores, Teliospores, and Basidiospores. Like P. monoica, outcrossing between mating types depends on a dispersal mechanism by an insect pollinator. U. pisi alternates between two plant hosts to proceed through its life cycle. E. cyparissias hosts the fungal spermatia and aeciospores, while a species in the family Fabaceae is host to the other spore types.[8]

Ecology

[edit]

Pseudoflower-insect interactions

[edit]

Rust fungi reproduction depends on insect visitation, analogous to pollination in flowers. Insects transfer spermatia (gametes) between different mating types of infected plants. The presence of the pseudoflower in plant communities may have consequences other than enhance the reproduction of the fungus. Studies in P. monoica demonstrated that insect visitation occurs more often when plants present a mixture of both pseudoflowers and natural occurring flowers, rather than when they occur alone in the plant. Combination of real flowers and pseudoflowers in plants increase the floral display through and apparent higher density, it also produces complementary attraction cues and complementary pollinators rewards, thus it has been proposed that they play a role in the pollination dynamics between insect and plants.[9] In U. pisi, insect pollinator visits are also necessary in order for two mating types of U. pisi to interact and result in fertilization.[3]

Pseudoflowers are visited by halictid, andrenic, bumble and oligolectic bees, muscid flies, and ants.[10]

See also

[edit]

Rust (fungus)

Parasitism

Mimicry in plants

References

[edit]
  1. ^ a b http://website.nbm-mnb.ca/mycologywebpages/NaturalHistoryOfFungi/Pucciniales.html
  2. ^ a b Roy, B. A. "Floral Mimicry by a Plant Pathogen." Nature 362 (1993): 56 - 58.
  3. ^ a b Pfunder, Monika, and Bitty A. Roy. "Pollinator-mediated interactions between a pathogenic fungus, Uromyces pisi (Pucciniaceae), and its host plant, Euphorbia cyparissias (Euphorbiaceae)." American Journal of Botany 87 (2000): 48 - 55.
  4. ^ Pfunder, Monika; Schürch, Stéphanie; Roy, Barbara A. "Sequence variation and geographic distribution of pseudoflower-forming rust fungi (Uromyces pisi s. lat.) on Euphorbia cyparissias". Mycological Research. 105 (1): 57–66. doi:10.1017/s0953756200003208.
  5. ^ a b c d Raguso, Robert A., and Barbara A. Roy. "‘Floral’scent production by Puccinia rust fungi that mimic flowers." Molecular Ecology 7.9 (1998): 1127-1136.
  6. ^ a b Kaiser, Roman (2006-02-10). "Flowers and Fungi Use Scents to Mimic Each Other". Science. 311 (5762): 806–807. doi:10.1126/science.1119499. ISSN 0036-8075. PMID 16469916.
  7. ^ Barilli E., Sillero J.C., Prats E., Rubiales D. "Resistance to rusts (Uromyces pisi and U. viciae-fabae) in pea." Czech J. Genet. Plant Breed 50 (2014): 135–143.
  8. ^ GAUMANN, E. Die Rostpilze Mitteleuropas. Beitrage zur Kryp-togamenflora der Schweiz Band 12 (1959): 1–1407.
  9. ^ Roy, B. A. (1994-03-01). "The Effects of Pathogen-Induced Pseudoflowers and Buttercups on Each Other's Insect Visitation". Ecology. 75 (2): 352–358. doi:10.2307/1939539. ISSN 1939-9170.
  10. ^ Roy, B.A.Raguso, R.A.Olfactory versus visual cues in a floral mimicry systemOecologia109 1997414426