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== Nuclear Movement in Response to Stimuli ==
= The Role of the Cytoskeleton in Plant Signaling and Behavior =
The plant [[cytoskeleton]] plays key roles in plant cellular signaling and mediates key processes in plant cellular responses underlying plant behavior and physiology. These processes include nuclear and chloroplastic movements in response to [[Biotic component|biotic]] and [[Abiotic component|abiotic]] stimuli, polar growth, and cell patterning. Plant cells face unique problems compared to [[animal cells]], which is reflected in the different characteristics and roles of the plant cytoskeleton.

== Characteristics of the Plant Cytoskeleton ==
Plant cells use [[actin]] as the major cytoskeletal component mediating vesicle trafficking, as opposed to animal cells, which primarily rely on [[Microtubule|microtubules]] to make up [[Endomembrane system|endomembrane]] trafficking networks<ref>{{Cite journal|last=Kost|first=Benedikt|last2=Chua|first2=Nam-Hai|date=2002-01-11|title=The Plant Cytoskeleton: Vacuoles and Cell Walls Make the Difference|url=https://www.cell.com/cell/abstract/S0092-8674(01)00634-1|journal=Cell|language=English|volume=108|issue=1|pages=9–12|doi=10.1016/S0092-8674(01)00634-1|issn=0092-8674|pmid=11792316}}</ref>. Microtubules in plants serve as constraining factors in polarized cell growth<ref>{{Cite journal|last=Kropf|first=Darryl L|last2=Bisgrove|first2=Sherryl R|last3=Hable|first3=Whitney E|date=1998-02|title=Cytoskeletal control of polar growth in plant cells|url=https://linkinghub.elsevier.com/retrieve/pii/S095506749880094X|journal=Current Opinion in Cell Biology|language=en|volume=10|issue=1|pages=117–122|doi=10.1016/S0955-0674(98)80094-X}}</ref>, and also are important during cell division, similarly to animals. The third major component of the cytoskeleton in animals cells, [[Intermediate filament|intermediate filaments]], has remained elusive in plants, with only a few studies suggesting the existence of intermediate filament-like proteins in plant cells<ref>{{Cite journal|last=Yang|first=C.|last2=Xing|first2=L.|last3=Zhai|first3=Z.|date=1992-03|title=Intermediate filaments in higher plant cells and their assembly in a cell-free system|url=http://link.springer.com/10.1007/BF01379279|journal=Protoplasma|language=en|volume=171|issue=1-2|pages=44–54|doi=10.1007/BF01379279|issn=0033-183X}}</ref>.

Plant cells are characterized by large vacuoles with narrow of regions of cytoplasm<ref>{{Cite journal|last=Marty|first=Francis|date=1999-04-01|title=Plant Vacuoles|url=http://www.plantcell.org/content/11/4/587|journal=The Plant Cell|language=en|volume=11|issue=4|pages=587–599|doi=10.1105/tpc.11.4.587|issn=1040-4651|pmc=PMC144210|pmid=10213780}}</ref>. This poses a kinetic problem to the movement of materials within the cell, as moving from one side of the cell would take an extremely long time without directed movements along the cytoskeleton.  Additionally, because of the rigid cell wall constraining cell shape, cell division and polarized growth necessitates tightly controlled spatial regulation. Plants use the dynamic nature cytoskeletal network to solve these problems.

== Cell Biology, Physiology, and Behavior ==
Contrary to the static nature often attributed to plants, plants are highly dynamic organisms. This includes processes at the cellular level, especially developmental processes underlying plant morphology and organelle organization in the cell, which both are mediated by the cytoskeleton. Cellular processes mediated by the plant cytoskeleton underly many aspects of plant physiology in response to biotic factors, such as pathogens, as well as abiotic factors, such as directional light or mechanical stimulation.

=== Nuclear Movement in Response to Stimuli ===
An important aspect of plant behavior includes responding to directional stimuli, which requires changes in cellular signaling to control spatial elements. The integration of stimuli in plant cells is not fully understood, but the movement of the cell nucleus provides one example of a cellular process that underlies plant behavior, and highlights the importance of the cytoskeleton in solving spatial problems within the plant cell. Unlike the static nature typically depicted in textbooks, the plant cell [[Cell nucleus|nucleus]] is a highly dynamic structure, constantly moving around cell via [[actin]] networks and [[Myosin|myosins]]<ref>{{Cite journal|last=Ketelaar|first=Tijs|last2=Faivre-Moskalenko|first2=Cendrine|last3=Esseling|first3=John J.|last4=de Ruijter|first4=Norbert C. A.|last5=Grierson|first5=Claire S.|last6=Dogterom|first6=Marileen|last7=Emons|first7=Anne Mie C.|date=2002-11|title=Positioning of Nuclei in Arabidopsis Root Hairs: An Actin-Regulated Process of Tip Growth|url=http://www.plantcell.org/lookup/doi/10.1105/tpc.005892|journal=The Plant Cell|language=en|volume=14|issue=11|pages=2941–2955|doi=10.1105/tpc.005892|issn=1040-4651|pmc=PMC152738|pmid=12417712}}</ref>. The nucleus undergoes a characteristic program during cell division to guide asymmetric cell division<ref>{{Cite journal|last=Smith|first=Laurie G.|date=2001-01|title=Plant cell division: building walls in the right places|url=http://www.nature.com/articles/35048050|journal=Nature Reviews Molecular Cell Biology|language=en|volume=2|issue=1|pages=33–39|doi=10.1038/35048050|issn=1471-0072}}</ref>, but there are several stimuli that have been demonstrated to cause movements of the nucleus in the plant cell<ref>{{Cite journal|last=Griffis|first=Anna Hare Newman|last2=Groves|first2=Norman Reid|last3=Zhou|first3=Xiao|last4=Meier|first4=Iris|date=2014|title=Nuclei in motion: movement and positioning of plant nuclei in development, signaling, symbiosis, and disease|url=https://www.frontiersin.org/articles/10.3389/fpls.2014.00129/full|journal=Frontiers in Plant Science|language=English|volume=5|doi=10.3389/fpls.2014.00129|issn=1664-462X|pmc=PMC3982112|pmid=24772115}}</ref>.
An important aspect of plant behavior includes responding to directional stimuli, which requires changes in cellular signaling to control spatial elements. The integration of stimuli in plant cells is not fully understood, but the movement of the cell nucleus provides one example of a cellular process that underlies plant behavior, and highlights the importance of the cytoskeleton in solving spatial problems within the plant cell. Unlike the static nature typically depicted in textbooks, the plant cell [[Cell nucleus|nucleus]] is a highly dynamic structure, constantly moving around cell via [[actin]] networks and [[Myosin|myosins]]<ref>{{Cite journal|last=Ketelaar|first=Tijs|last2=Faivre-Moskalenko|first2=Cendrine|last3=Esseling|first3=John J.|last4=de Ruijter|first4=Norbert C. A.|last5=Grierson|first5=Claire S.|last6=Dogterom|first6=Marileen|last7=Emons|first7=Anne Mie C.|date=2002-11|title=Positioning of Nuclei in Arabidopsis Root Hairs: An Actin-Regulated Process of Tip Growth|url=http://www.plantcell.org/lookup/doi/10.1105/tpc.005892|journal=The Plant Cell|language=en|volume=14|issue=11|pages=2941–2955|doi=10.1105/tpc.005892|issn=1040-4651|pmc=PMC152738|pmid=12417712}}</ref>. The nucleus undergoes a characteristic program during cell division to guide asymmetric cell division<ref>{{Cite journal|last=Smith|first=Laurie G.|date=2001-01|title=Plant cell division: building walls in the right places|url=http://www.nature.com/articles/35048050|journal=Nature Reviews Molecular Cell Biology|language=en|volume=2|issue=1|pages=33–39|doi=10.1038/35048050|issn=1471-0072}}</ref>, but there are several stimuli that have been demonstrated to cause movements of the nucleus in the plant cell<ref>{{Cite journal|last=Griffis|first=Anna Hare Newman|last2=Groves|first2=Norman Reid|last3=Zhou|first3=Xiao|last4=Meier|first4=Iris|date=2014|title=Nuclei in motion: movement and positioning of plant nuclei in development, signaling, symbiosis, and disease|url=https://www.frontiersin.org/articles/10.3389/fpls.2014.00129/full|journal=Frontiers in Plant Science|language=English|volume=5|doi=10.3389/fpls.2014.00129|issn=1664-462X|pmc=PMC3982112|pmid=24772115}}</ref>.


=== Blue Light ===
A well-studied stimulus is strong blue light, which drives movement of nuclei to anticlinal (perpendicular to the plane of the leaf) [[Cell wall|cell walls]] in [[Mesophyll|mesophyl]]<nowiki/>l and epidermal cells of ''[[Arabidopsis thaliana]]'' plants.<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Sakai|first2=Tatsuya|last3=Takagi|first3=Shingo|date=2007-09-01|title=Blue Light-Dependent Nuclear Positioning in Arabidopsis thaliana Leaf Cells|url=https://academic.oup.com/pcp/article/48/9/1291/1814996|journal=Plant and Cell Physiology|language=en|volume=48|issue=9|pages=1291–1298|doi=10.1093/pcp/pcm095|issn=0032-0781}}</ref> Chloroplasts moving in response to blue light associate with the nucleus to move the nucleus to the appropriate location<ref>{{Cite journal|last=Higa|first=Takeshi|last2=Suetsugu|first2=Noriyuki|last3=Kong|first3=Sam-Geun|last4=Wada|first4=Masamitsu|date=2014-03-18|title=Actin-dependent plastid movement is required for motive force generation in directional nuclear movement in plants|url=https://www.pnas.org/content/111/11/4327|journal=Proceedings of the National Academy of Sciences|language=en|volume=111|issue=11|pages=4327–4331|doi=10.1073/pnas.1317902111|issn=0027-8424|pmc=PMC3964046|pmid=24591587}}</ref>. This is highly dependent on the blue light receptor [[phototropin]] and the actin cytoskeleton, as actin bundles are seen to form along the anticlinal wall in blue light<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Minamino|first2=Ryoko|last3=Takagi|first3=Shingo|date=2010-03-01|title=Actin Reorganization Underlies Phototropin-Dependent Positioning of Nuclei in Arabidopsis Leaf Cells|url=http://www.plantphysiol.org/content/152/3/1309|journal=Plant Physiology|language=en|volume=152|issue=3|pages=1309–1319|doi=10.1104/pp.109.149526|issn=0032-0889|pmc=PMC2832274|pmid=20107027}}</ref>. A protein called ANGUSTIFOLIA was also recently discovered to regulate nucleus movement in the dark by forming a complex that adjusts the alignment of actin filaments<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Ohnishi|first2=Haruna|last3=Tamura|first3=Kentaro|last4=Fukao|first4=Yoichiro|last5=Furuya|first5=Tomoyuki|last6=Hattori|first6=Koro|last7=Tsukaya|first7=Hirokazu|last8=Hara-Nishimura|first8=Ikuko|date=2019-01-01|title=ANGUSTIFOLIA Regulates Actin Filament Alignment for Nuclear Positioning in Leaves|url=http://www.plantphysiol.org/content/179/1/233|journal=Plant Physiology|language=en|volume=179|issue=1|pages=233–247|doi=10.1104/pp.18.01150|issn=0032-0889|pmc=PMC6324246|pmid=30404821}}</ref>. The movement of the nucleus in response to blue light may serve several physiological purposes<ref name=":0">{{Cite journal|last=Iwabuchi|first=Kosei|last2=Takagi|first2=Shingo|date=2008-04-01|title=How and why do plant nuclei move in response to light?|url=https://doi.org/10.4161/psb.3.4.5213|journal=Plant Signaling & Behavior|volume=3|issue=4|pages=266–268|doi=10.4161/psb.3.4.5213|pmc=PMC2634198|pmid=19704650}}</ref>. The first is to avoid damaging mutations caused by [[Ultraviolet|UV]] radiation, as the nucleus stores the genetic material of a cell. A key problem faced as photosynthetic organisms transitioned from ocean to land was avoiding excessive mutations caused by UV radiation, but by moving the nucleus in response to light, damage caused by UV light could be limited. Another purpose may be to localize the nucleus near key receptors, such as [[Phytochrome|phytochrome,]] to facilitate spatial integration and transduction of cellular signals into the nucleus, especially when considering the necessity of phytochrome import into the nucleus for changes in gene expression in response to red light.<ref name=":0" />
A well-studied stimulus is strong blue light, which drives movement of nuclei to anticlinal (perpendicular to the plane of the leaf) [[Cell wall|cell walls]] in [[Mesophyll|mesophyl]]<nowiki/>l and epidermal cells of ''[[Arabidopsis thaliana]]'' plants.<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Sakai|first2=Tatsuya|last3=Takagi|first3=Shingo|date=2007-09-01|title=Blue Light-Dependent Nuclear Positioning in Arabidopsis thaliana Leaf Cells|url=https://academic.oup.com/pcp/article/48/9/1291/1814996|journal=Plant and Cell Physiology|language=en|volume=48|issue=9|pages=1291–1298|doi=10.1093/pcp/pcm095|issn=0032-0781}}</ref> Chloroplasts moving in response to blue light associate with the nucleus to move the nucleus to the appropriate location<ref>{{Cite journal|last=Higa|first=Takeshi|last2=Suetsugu|first2=Noriyuki|last3=Kong|first3=Sam-Geun|last4=Wada|first4=Masamitsu|date=2014-03-18|title=Actin-dependent plastid movement is required for motive force generation in directional nuclear movement in plants|url=https://www.pnas.org/content/111/11/4327|journal=Proceedings of the National Academy of Sciences|language=en|volume=111|issue=11|pages=4327–4331|doi=10.1073/pnas.1317902111|issn=0027-8424|pmc=PMC3964046|pmid=24591587}}</ref>. This is highly dependent on the blue light receptor [[phototropin]] and the actin cytoskeleton, as actin bundles are seen to form along the anticlinal wall in blue light<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Minamino|first2=Ryoko|last3=Takagi|first3=Shingo|date=2010-03-01|title=Actin Reorganization Underlies Phototropin-Dependent Positioning of Nuclei in Arabidopsis Leaf Cells|url=http://www.plantphysiol.org/content/152/3/1309|journal=Plant Physiology|language=en|volume=152|issue=3|pages=1309–1319|doi=10.1104/pp.109.149526|issn=0032-0889|pmc=PMC2832274|pmid=20107027}}</ref>. A protein called ANGUSTIFOLIA was also recently discovered to regulate nucleus movement in the dark by forming a complex that adjusts the alignment of actin filaments<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Ohnishi|first2=Haruna|last3=Tamura|first3=Kentaro|last4=Fukao|first4=Yoichiro|last5=Furuya|first5=Tomoyuki|last6=Hattori|first6=Koro|last7=Tsukaya|first7=Hirokazu|last8=Hara-Nishimura|first8=Ikuko|date=2019-01-01|title=ANGUSTIFOLIA Regulates Actin Filament Alignment for Nuclear Positioning in Leaves|url=http://www.plantphysiol.org/content/179/1/233|journal=Plant Physiology|language=en|volume=179|issue=1|pages=233–247|doi=10.1104/pp.18.01150|issn=0032-0889|pmc=PMC6324246|pmid=30404821}}</ref>. The movement of the nucleus in response to blue light may serve several physiological purposes<ref name=":0">{{Cite journal|last=Iwabuchi|first=Kosei|last2=Takagi|first2=Shingo|date=2008-04-01|title=How and why do plant nuclei move in response to light?|url=https://doi.org/10.4161/psb.3.4.5213|journal=Plant Signaling & Behavior|volume=3|issue=4|pages=266–268|doi=10.4161/psb.3.4.5213|pmc=PMC2634198|pmid=19704650}}</ref>. The first is to avoid damaging mutations caused by [[Ultraviolet|UV]] radiation, as the nucleus stores the genetic material of a cell. A key problem faced as photosynthetic organisms transitioned from ocean to land was avoiding excessive mutations caused by UV radiation, but by moving the nucleus in response to light, damage caused by UV light could be limited. Another purpose may be to localize the nucleus near key receptors, such as [[Phytochrome|phytochrome,]] to facilitate spatial integration and transduction of cellular signals into the nucleus, especially when considering the necessity of phytochrome import into the nucleus for changes in gene expression in response to red light.<ref name=":0" />

=== Mechanical Stimulation ===


Nuclear movement also occurs in response to mechanical stimulation. The nuclei of cultured [[ovule]] [[Parenchyma (botany)|parenchyma]] [[Nicotiana tabacum|tobacco]] cells were found to move directly to the site of probing by a fine glass pipette via cytoplasmic strands<ref>{{Cite journal|last=Qu|first=Liang-Huan|last2=Sun|first2=Meng-Xiang|date=2008-09|title=Cytoplasmic compartmental response to local mechanical stimulation of internal tissue cells|url=http://link.springer.com/10.1007/s00709-008-0304-0|journal=Protoplasma|language=en|volume=233|issue=1-2|pages=51–59|doi=10.1007/s00709-008-0304-0|issn=0033-183X}}</ref>, which contain actin filaments specialized to carry out cytoplasmic streaming. This is likely a response co-opted from cytoplasmic streaming, but a receptor or other downstream signaling components underlying this cellular response have not been identified. Nonetheless mechanical stimulation is a potent signal resulting in nuclear movement, and suggests that nuclear movement may be a process important for integration of mechanical stimulation during [[thigmotropism]], [[gravitropism]], or cellular interactions during development<ref>{{Cite journal|last=Qu|first=Liang-Huan|last2=Sun|first2=Meng-Xiang|date=2007-08-01|title=The plant cell nucleus is constantly alert and highly sensitive to repetitive local mechanical stimulations|url=https://doi.org/10.1007/s00299-007-0343-6|journal=Plant Cell Reports|language=en|volume=26|issue=8|pages=1187–1193|doi=10.1007/s00299-007-0343-6|issn=1432-203X}}</ref>.
Nuclear movement also occurs in response to mechanical stimulation. The nuclei of cultured [[ovule]] [[Parenchyma (botany)|parenchyma]] [[Nicotiana tabacum|tobacco]] cells were found to move directly to the site of probing by a fine glass pipette via cytoplasmic strands<ref>{{Cite journal|last=Qu|first=Liang-Huan|last2=Sun|first2=Meng-Xiang|date=2008-09|title=Cytoplasmic compartmental response to local mechanical stimulation of internal tissue cells|url=http://link.springer.com/10.1007/s00709-008-0304-0|journal=Protoplasma|language=en|volume=233|issue=1-2|pages=51–59|doi=10.1007/s00709-008-0304-0|issn=0033-183X}}</ref>, which contain actin filaments specialized to carry out cytoplasmic streaming. This is likely a response co-opted from cytoplasmic streaming, but a receptor or other downstream signaling components underlying this cellular response have not been identified. Nonetheless mechanical stimulation is a potent signal resulting in nuclear movement, and suggests that nuclear movement may be a process important for integration of mechanical stimulation during [[thigmotropism]], [[gravitropism]], or cellular interactions during development<ref>{{Cite journal|last=Qu|first=Liang-Huan|last2=Sun|first2=Meng-Xiang|date=2007-08-01|title=The plant cell nucleus is constantly alert and highly sensitive to repetitive local mechanical stimulations|url=https://doi.org/10.1007/s00299-007-0343-6|journal=Plant Cell Reports|language=en|volume=26|issue=8|pages=1187–1193|doi=10.1007/s00299-007-0343-6|issn=1432-203X}}</ref>.


=== Symbionts ===
Recognition of microbial organisms also results in nuclear movement. During colonization by beneficial [[rhizobia]], which begins at the root hair tip, the nucleus moves to the site of colonization and guides the formation and direction of movement of the infection thread, a structure that houses the colonizing rhizobium<ref>{{Cite journal|last=FÅHRAEUS|first=GÖSTA|date=1957|title=The Infection of Clover Root Hairs by Nodule Bacteria Studied by a Simple Glass Slide Technique|url=https://www.microbiologyresearch.org/content/journal/micro/10.1099/00221287-16-2-374|journal=Microbiology,|volume=16|issue=2|pages=374–381|doi=10.1099/00221287-16-2-374|issn=1350-0872}}</ref>. This requires large scale cytoskeletal rearrangment, as well as cytoskeleton-mediated movement of the nucleus<ref>{{Cite journal|last=Tsyganov|first=Viktor E.|last2=Kitaeva|first2=Anna B.|last3=Demchenko|first3=Kirill N.|date=2019-12-13|title=Comparative analysis of tubulin cytoskeleton rearrangements in nodules of
Recognition of microbial organisms also results in nuclear movement. During colonization by beneficial [[rhizobia]], which begins at the root hair tip, the nucleus moves to the site of colonization and guides the formation and direction of movement of the infection thread, a structure that houses the colonizing rhizobium<ref>{{Cite journal|last=FÅHRAEUS|first=GÖSTA|date=1957|title=The Infection of Clover Root Hairs by Nodule Bacteria Studied by a Simple Glass Slide Technique|url=https://www.microbiologyresearch.org/content/journal/micro/10.1099/00221287-16-2-374|journal=Microbiology,|volume=16|issue=2|pages=374–381|doi=10.1099/00221287-16-2-374|issn=1350-0872}}</ref>. This requires large scale cytoskeletal rearrangment, as well as cytoskeleton-mediated movement of the nucleus<ref>{{Cite journal|last=Tsyganov|first=Viktor E.|last2=Kitaeva|first2=Anna B.|last3=Demchenko|first3=Kirill N.|date=2019-12-13|title=Comparative analysis of tubulin cytoskeleton rearrangements in nodules of
Medicago truncatula
Medicago truncatula
Line 22: Line 15:
Pisum sativum|url=http://dx.doi.org/10.1002/9781119409144.ch68|journal=The Model Legume Medicago truncatula|pages=547–543|doi=10.1002/9781119409144.ch68}}</ref>. Similarly, [[Arbuscular mycorrhiza|arbuscular mycorrhizae]] symbiosis involves extensive nuclear movement, which apears to guide formation of microtubule structures that steers penetration by the fungal [[hypha]]<ref>{{Cite journal|last=Genre|first=Andrea|last2=Chabaud|first2=Mireille|last3=Timmers|first3=Ton|last4=Bonfante|first4=Paola|last5=Barker|first5=David G.|date=2005-12|title=Arbuscular Mycorrhizal Fungi Elicit a Novel Intracellular Apparatus in Medicago truncatula Root Epidermal Cells before Infection|url=http://www.plantcell.org/lookup/doi/10.1105/tpc.105.035410|journal=The Plant Cell|language=en|volume=17|issue=12|pages=3489–3499|doi=10.1105/tpc.105.035410|issn=1040-4651|pmc=PMC1315383|pmid=16284314}}</ref>.
Pisum sativum|url=http://dx.doi.org/10.1002/9781119409144.ch68|journal=The Model Legume Medicago truncatula|pages=547–543|doi=10.1002/9781119409144.ch68}}</ref>. Similarly, [[Arbuscular mycorrhiza|arbuscular mycorrhizae]] symbiosis involves extensive nuclear movement, which apears to guide formation of microtubule structures that steers penetration by the fungal [[hypha]]<ref>{{Cite journal|last=Genre|first=Andrea|last2=Chabaud|first2=Mireille|last3=Timmers|first3=Ton|last4=Bonfante|first4=Paola|last5=Barker|first5=David G.|date=2005-12|title=Arbuscular Mycorrhizal Fungi Elicit a Novel Intracellular Apparatus in Medicago truncatula Root Epidermal Cells before Infection|url=http://www.plantcell.org/lookup/doi/10.1105/tpc.105.035410|journal=The Plant Cell|language=en|volume=17|issue=12|pages=3489–3499|doi=10.1105/tpc.105.035410|issn=1040-4651|pmc=PMC1315383|pmid=16284314}}</ref>.


=== Pathogens ===
Importantly, cytoskeleton-mediated nuclear movement is critical for response of plants to pathogenic microorganisms. This is best studied in [[Oomycete|oomycetes]], a devastating pathogenic organism. In [[Potato|potato cells]], oomycete contact results in rapid movement of the nucleus to the site of contact, which initiates rapid deposition of cell wall material and restructuring of the cytoplasmic elements<ref>{{Cite journal|last=Freytag|first=Sibylle|last2=Arabatzis|first2=Nikolaos|last3=Hahlbrock|first3=Klaus|last4=Schmelzer|first4=Elmon|date=1994-06|title=Reversible cytoplasmic rearrangements precede wall apposition, hypersensitive cell death and defense-related gene activation in potato/Phytophthora infestans interactions|url=http://link.springer.com/10.1007/BF00201043|journal=Planta|language=en|volume=194|issue=1|doi=10.1007/BF00201043|issn=0032-0935}}</ref>. This can block invasion by the oomycete, or if the oomycete successfully penetrates the cell, can initiate a [[hypersensitive response]], killing the cell and preventing further propagation of the pathogen. However, in plants that are not resistant to oomycete infection, the nucleus does not move to the site of oomycete contact, and the oomycete proceeds to devestate the plant, indicating the importance of nuclear transport for resistance against oomycete pathogens.
Importantly, cytoskeleton-mediated nuclear movement is critical for response of plants to pathogenic microorganisms. This is best studied in [[Oomycete|oomycetes]], a devastating pathogenic organism. In [[Potato|potato cells]], oomycete contact results in rapid movement of the nucleus to the site of contact, which initiates rapid deposition of cell wall material and restructuring of the cytoplasmic elements<ref>{{Cite journal|last=Freytag|first=Sibylle|last2=Arabatzis|first2=Nikolaos|last3=Hahlbrock|first3=Klaus|last4=Schmelzer|first4=Elmon|date=1994-06|title=Reversible cytoplasmic rearrangements precede wall apposition, hypersensitive cell death and defense-related gene activation in potato/Phytophthora infestans interactions|url=http://link.springer.com/10.1007/BF00201043|journal=Planta|language=en|volume=194|issue=1|doi=10.1007/BF00201043|issn=0032-0935}}</ref>. This can block invasion by the oomycete, or if the oomycete successfully penetrates the cell, can initiate a [[hypersensitive response]], killing the cell and preventing further propagation of the pathogen. However, in plants that are not resistant to oomycete infection, the nucleus does not move to the site of oomycete contact, and the oomycete proceeds to devestate the plant, indicating the importance of nuclear transport for resistance against oomycete pathogens.


These examples of nuclear movement in response to biotic and abiotic stimuli highlight the role of the nucleus as a highly mobile command center necessary for integration of cell signaling and also emphasize the importance of cytoskeletal structure in mediating the transduction of signaling from outside the cell to the nucleus. However there is still a great deal left unknown in how exactly an extracellular stimulus leads to cytoskeletal rearrangement, nuclear movement, and ultimately integration of stimuli to guide plant behavior.
These examples of nuclear movement in response to biotic and abiotic stimuli highlight the role of the nucleus as a highly mobile command center necessary for integration of cell signaling and also emphasize the importance of cytoskeletal structure in mediating the transduction of signaling from outside the cell to the nucleus. However there is still a great deal left unknown in how exactly an extracellular stimulus leads to cytoskeletal rearrangement, nuclear movement, and ultimately integration of stimuli to guide plant behavior.

=== Other Biological Processes ===

* Microtubule rearrangement in response to ethylene during [[photomorphogenesis]] results in lateral cell expansion that drive shoot thickening<ref>{{Cite journal|last=Le|first=Jie|last2=Vandenbussche|first2=Filip|last3=De Cnodder|first3=Tinne|last4=Van Der Straeten|first4=Dominique|last5=Verbelen|first5=Jean-Pierre|date=2005-09|title=Cell Elongation and Microtubule Behavior in the Arabidopsis Hypocotyl: Responses to Ethylene and Auxin|url=http://link.springer.com/10.1007/s00344-005-0044-8|journal=Journal of Plant Growth Regulation|language=en|volume=24|issue=3|pages=166–178|doi=10.1007/s00344-005-0044-8|issn=0721-7595}}</ref>.
* Polar auxin transport depends on actin dynamics<ref>{{Cite journal|last=Zou|first=Minxia|last2=Ren|first2=Haiyun|last3=Li|first3=Jiejie|date=2019-09-01|title=An Auxin Transport Inhibitor Targets Villin-Mediated Actin Dynamics to Regulate Polar Auxin Transport|url=http://www.plantphysiol.org/content/181/1/161|journal=Plant Physiology|language=en|volume=181|issue=1|pages=161–178|doi=10.1104/pp.19.00064|issn=0032-0889|pmc=PMC6716258|pmid=31311831}}</ref>.
*Polar Cell growth<ref>{{Cite journal|last=Kropf|first=Darryl L|last2=Bisgrove|first2=Sherryl R|last3=Hable|first3=Whitney E|date=1998-02|title=Cytoskeletal control of polar growth in plant cells|url=http://dx.doi.org/10.1016/s0955-0674(98)80094-x|journal=Current Opinion in Cell Biology|volume=10|issue=1|pages=117–122|doi=10.1016/s0955-0674(98)80094-x|issn=0955-0674}}</ref>
*Movement of chloroplasts to avoid photooxidative damage<ref>{{Cite journal|last=Iwabuchi|first=Kosei|last2=Takagi|first2=Shingo|date=2010-08-01|title=Actin-based mechanisms for light-dependent intracellular positioning of nuclei and chloroplasts in Arabidopsis|url=https://doi.org/10.4161/psb.5.8.12233|journal=Plant Signaling & Behavior|volume=5|issue=8|pages=1010–1013|doi=10.4161/psb.5.8.12233|pmc=PMC3115182|pmid=20724834}}</ref><ref>{{Cite journal|last=Anielska-Mazur|first=Anna|last2=Bernaś|first2=Tytus|last3=Gabryś|first3=Halina|date=2009|title=In vivo reorganization of the actin cytoskeleton in leaves of Nicotiana tabacum L. transformed with plastin-GFP. Correlation with light-activated chloroplast responses|url=http://dx.doi.org/10.1186/1471-2229-9-64|journal=BMC Plant Biology|volume=9|issue=1|pages=64|doi=10.1186/1471-2229-9-64|issn=1471-2229}}</ref>.


== References ==
== References ==

Revision as of 02:45, 4 June 2020

Nuclear Movement in Response to Stimuli

An important aspect of plant behavior includes responding to directional stimuli, which requires changes in cellular signaling to control spatial elements. The integration of stimuli in plant cells is not fully understood, but the movement of the cell nucleus provides one example of a cellular process that underlies plant behavior, and highlights the importance of the cytoskeleton in solving spatial problems within the plant cell. Unlike the static nature typically depicted in textbooks, the plant cell nucleus is a highly dynamic structure, constantly moving around cell via actin networks and myosins[1]. The nucleus undergoes a characteristic program during cell division to guide asymmetric cell division[2], but there are several stimuli that have been demonstrated to cause movements of the nucleus in the plant cell[3].

Blue Light

A well-studied stimulus is strong blue light, which drives movement of nuclei to anticlinal (perpendicular to the plane of the leaf) cell walls in mesophyll and epidermal cells of Arabidopsis thaliana plants.[4] Chloroplasts moving in response to blue light associate with the nucleus to move the nucleus to the appropriate location[5]. This is highly dependent on the blue light receptor phototropin and the actin cytoskeleton, as actin bundles are seen to form along the anticlinal wall in blue light[6]. A protein called ANGUSTIFOLIA was also recently discovered to regulate nucleus movement in the dark by forming a complex that adjusts the alignment of actin filaments[7]. The movement of the nucleus in response to blue light may serve several physiological purposes[8]. The first is to avoid damaging mutations caused by UV radiation, as the nucleus stores the genetic material of a cell. A key problem faced as photosynthetic organisms transitioned from ocean to land was avoiding excessive mutations caused by UV radiation, but by moving the nucleus in response to light, damage caused by UV light could be limited. Another purpose may be to localize the nucleus near key receptors, such as phytochrome, to facilitate spatial integration and transduction of cellular signals into the nucleus, especially when considering the necessity of phytochrome import into the nucleus for changes in gene expression in response to red light.[8]

Mechanical Stimulation

Nuclear movement also occurs in response to mechanical stimulation. The nuclei of cultured ovule parenchyma tobacco cells were found to move directly to the site of probing by a fine glass pipette via cytoplasmic strands[9], which contain actin filaments specialized to carry out cytoplasmic streaming. This is likely a response co-opted from cytoplasmic streaming, but a receptor or other downstream signaling components underlying this cellular response have not been identified. Nonetheless mechanical stimulation is a potent signal resulting in nuclear movement, and suggests that nuclear movement may be a process important for integration of mechanical stimulation during thigmotropism, gravitropism, or cellular interactions during development[10].

Symbionts

Recognition of microbial organisms also results in nuclear movement. During colonization by beneficial rhizobia, which begins at the root hair tip, the nucleus moves to the site of colonization and guides the formation and direction of movement of the infection thread, a structure that houses the colonizing rhizobium[11]. This requires large scale cytoskeletal rearrangment, as well as cytoskeleton-mediated movement of the nucleus[12]. Similarly, arbuscular mycorrhizae symbiosis involves extensive nuclear movement, which apears to guide formation of microtubule structures that steers penetration by the fungal hypha[13].

Pathogens

Importantly, cytoskeleton-mediated nuclear movement is critical for response of plants to pathogenic microorganisms. This is best studied in oomycetes, a devastating pathogenic organism. In potato cells, oomycete contact results in rapid movement of the nucleus to the site of contact, which initiates rapid deposition of cell wall material and restructuring of the cytoplasmic elements[14]. This can block invasion by the oomycete, or if the oomycete successfully penetrates the cell, can initiate a hypersensitive response, killing the cell and preventing further propagation of the pathogen. However, in plants that are not resistant to oomycete infection, the nucleus does not move to the site of oomycete contact, and the oomycete proceeds to devestate the plant, indicating the importance of nuclear transport for resistance against oomycete pathogens.

These examples of nuclear movement in response to biotic and abiotic stimuli highlight the role of the nucleus as a highly mobile command center necessary for integration of cell signaling and also emphasize the importance of cytoskeletal structure in mediating the transduction of signaling from outside the cell to the nucleus. However there is still a great deal left unknown in how exactly an extracellular stimulus leads to cytoskeletal rearrangement, nuclear movement, and ultimately integration of stimuli to guide plant behavior.

References

  1. ^ Ketelaar, Tijs; Faivre-Moskalenko, Cendrine; Esseling, John J.; de Ruijter, Norbert C. A.; Grierson, Claire S.; Dogterom, Marileen; Emons, Anne Mie C. (2002-11). "Positioning of Nuclei in Arabidopsis Root Hairs: An Actin-Regulated Process of Tip Growth". The Plant Cell. 14 (11): 2941–2955. doi:10.1105/tpc.005892. ISSN 1040-4651. PMC 152738. PMID 12417712. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
  2. ^ Smith, Laurie G. (2001-01). "Plant cell division: building walls in the right places". Nature Reviews Molecular Cell Biology. 2 (1): 33–39. doi:10.1038/35048050. ISSN 1471-0072. {{cite journal}}: Check date values in: |date= (help)
  3. ^ Griffis, Anna Hare Newman; Groves, Norman Reid; Zhou, Xiao; Meier, Iris (2014). "Nuclei in motion: movement and positioning of plant nuclei in development, signaling, symbiosis, and disease". Frontiers in Plant Science. 5. doi:10.3389/fpls.2014.00129. ISSN 1664-462X. PMC 3982112. PMID 24772115.{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)
  4. ^ Iwabuchi, Kosei; Sakai, Tatsuya; Takagi, Shingo (2007-09-01). "Blue Light-Dependent Nuclear Positioning in Arabidopsis thaliana Leaf Cells". Plant and Cell Physiology. 48 (9): 1291–1298. doi:10.1093/pcp/pcm095. ISSN 0032-0781.
  5. ^ Higa, Takeshi; Suetsugu, Noriyuki; Kong, Sam-Geun; Wada, Masamitsu (2014-03-18). "Actin-dependent plastid movement is required for motive force generation in directional nuclear movement in plants". Proceedings of the National Academy of Sciences. 111 (11): 4327–4331. doi:10.1073/pnas.1317902111. ISSN 0027-8424. PMC 3964046. PMID 24591587.{{cite journal}}: CS1 maint: PMC format (link)
  6. ^ Iwabuchi, Kosei; Minamino, Ryoko; Takagi, Shingo (2010-03-01). "Actin Reorganization Underlies Phototropin-Dependent Positioning of Nuclei in Arabidopsis Leaf Cells". Plant Physiology. 152 (3): 1309–1319. doi:10.1104/pp.109.149526. ISSN 0032-0889. PMC 2832274. PMID 20107027.{{cite journal}}: CS1 maint: PMC format (link)
  7. ^ Iwabuchi, Kosei; Ohnishi, Haruna; Tamura, Kentaro; Fukao, Yoichiro; Furuya, Tomoyuki; Hattori, Koro; Tsukaya, Hirokazu; Hara-Nishimura, Ikuko (2019-01-01). "ANGUSTIFOLIA Regulates Actin Filament Alignment for Nuclear Positioning in Leaves". Plant Physiology. 179 (1): 233–247. doi:10.1104/pp.18.01150. ISSN 0032-0889. PMC 6324246. PMID 30404821.{{cite journal}}: CS1 maint: PMC format (link)
  8. ^ a b Iwabuchi, Kosei; Takagi, Shingo (2008-04-01). "How and why do plant nuclei move in response to light?". Plant Signaling & Behavior. 3 (4): 266–268. doi:10.4161/psb.3.4.5213. PMC 2634198. PMID 19704650.{{cite journal}}: CS1 maint: PMC format (link)
  9. ^ Qu, Liang-Huan; Sun, Meng-Xiang (2008-09). "Cytoplasmic compartmental response to local mechanical stimulation of internal tissue cells". Protoplasma. 233 (1–2): 51–59. doi:10.1007/s00709-008-0304-0. ISSN 0033-183X. {{cite journal}}: Check date values in: |date= (help)
  10. ^ Qu, Liang-Huan; Sun, Meng-Xiang (2007-08-01). "The plant cell nucleus is constantly alert and highly sensitive to repetitive local mechanical stimulations". Plant Cell Reports. 26 (8): 1187–1193. doi:10.1007/s00299-007-0343-6. ISSN 1432-203X.
  11. ^ FÅHRAEUS, GÖSTA (1957). "The Infection of Clover Root Hairs by Nodule Bacteria Studied by a Simple Glass Slide Technique". Microbiology,. 16 (2): 374–381. doi:10.1099/00221287-16-2-374. ISSN 1350-0872.{{cite journal}}: CS1 maint: extra punctuation (link)
  12. ^ Tsyganov, Viktor E.; Kitaeva, Anna B.; Demchenko, Kirill N. (2019-12-13). "Comparative analysis of tubulin cytoskeleton rearrangements in nodules of Medicago truncatula and Pisum sativum". The Model Legume Medicago truncatula: 547–543. doi:10.1002/9781119409144.ch68. {{cite journal}}: line feed character in |title= at position 74 (help)
  13. ^ Genre, Andrea; Chabaud, Mireille; Timmers, Ton; Bonfante, Paola; Barker, David G. (2005-12). "Arbuscular Mycorrhizal Fungi Elicit a Novel Intracellular Apparatus in Medicago truncatula Root Epidermal Cells before Infection". The Plant Cell. 17 (12): 3489–3499. doi:10.1105/tpc.105.035410. ISSN 1040-4651. PMC 1315383. PMID 16284314. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
  14. ^ Freytag, Sibylle; Arabatzis, Nikolaos; Hahlbrock, Klaus; Schmelzer, Elmon (1994-06). "Reversible cytoplasmic rearrangements precede wall apposition, hypersensitive cell death and defense-related gene activation in potato/Phytophthora infestans interactions". Planta. 194 (1). doi:10.1007/BF00201043. ISSN 0032-0935. {{cite journal}}: Check date values in: |date= (help)