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[[File:Actinodaphne young leaf flush DSC1272.jpg|alt=|thumb|Leaf flush in an ''[[Actinodaphne]]'' tree showing young whitish leaves turning pale green against the darker green mature leaves. ]]
[[File:Actinodaphne young leaf flush DSC1272.jpg|alt=|thumb|A flush of ''[[Actinodaphne]]'' young leaves.]]'''Leaf flushing''' is the production of a flush of new leaves typically produced simultaneously on all branches of a bare plant or tree. Young leaves often have less [[chlorophyll]] and the leaf flush may be white or red, the latter due to presence of pigments, particularly [[anthocyanin]]s.<ref>{{Cite journal|last=Dominy|first=Nathaniel J.|last2=Lucas|first2=Peter W.|last3=Ramsden|first3=Lawrence W.|last4=Riba-Hernandez|first4=Pablo|last5=Stoner|first5=Kathryn E.|last6=Turner|first6=Ian M.|date=July 2002|title=Why are young leaves red?|url=http://doi.wiley.com/10.1034/j.1600-0706.2002.980117.x|journal=Oikos|language=en|volume=98|issue=1|pages=163–176|doi=10.1034/j.1600-0706.2002.980117.x}}</ref> Leaf flushing succeeds [[Abscission#In plants|leaf fall]], and is delayed by winter in the temperate zone or by extreme dryness in the tropics. Leaf fall and leaf flushing in tropical [[deciduous]] forests can overlap in some species, called ''leaf-exchanging'' species, producing new leaves during the same period when old leaves are shed or almost immediately after.<ref name="Elliott">{{cite journal|last1=Elliott|first1=Stephen|last2=Baker|first2=Patrick J.|last3=Borchert|first3=Rolf|title=Leaf flushing during the dry season: the paradox of Asian monsoon forests|journal=Global Ecology and Biogeography|date=May 2006|volume=15|issue=3|pages=248–257|doi=10.1111/j.1466-8238.2006.00213.x|url=http://ctfs.si.edu/Public/pdfs/ToDelete/Elliott_etal_2006_GlobalEcolBiogeogr.pdf}}</ref><ref>{{cite journal|last1=Heatwole|first1=Harold|last2=Lowman|first2=Margaret D.|last3=Donovan|first3=Christopher|last4=McCoy|first4=Maria|title=Phenology of leaf-flushing and macroarthropod abundances in canopies of ''Eucalyptus'' saplings|journal=Selbyana|date=1 January 1997|volume=18|issue=2|pages=200–214|jstor=41760435}}</ref> Leaf-flushing may be synchronized among trees of a single species or even across species in an area.<ref>{{cite journal|doi=10.2307/2388052 |jstor=2388052 |title=The Causes and Consequences of Synchronous Flushing in a Dry Tropical Forest |journal=Biotropica |volume=16 |issue=3 |pages=193 |year=1984 |last1=Lieberman |first1=Diana |last2=Lieberman |first2=Milton }}</ref> In the seasonal tropics, leaf flushing [[phenology]] may be influenced by herbivory and water stress.<ref>{{cite journal|last1=Murali|first1=K. S.|last2=Sukumar|first2=R.|title=Leaf flushing phenology and herbivory in a tropical dry deciduous forest, southern India|journal=Oecologia|date=1993|volume=94|issue=1|pages=114–119|url=http://www.geocities.ws/murali_kallur/oecologia.pdf|doi=10.1007/BF00317311|pmid=28313868}}</ref><ref>{{Cite journal|last=Elliott|first=Stephen|last2=Baker|first2=Patrick J.|last3=Borchert|first3=Rolf|date=2006|title=Leaf Flushing during the Dry Season: The Paradox of Asian Monsoon Forests|url=https://www.jstor.org/stable/3697587|journal=Global Ecology and Biogeography|volume=15|issue=3|pages=248–257|issn=1466-822X}}</ref>
[[File:Plant Canarium strictum tree with leaf flush DSCN0171.jpg|thumb|A ''[[Canarium strictum]]'' tree in red leaf flush.]]
'''Leaf flushing''' is the production of a flush of new leaves typically produced simultaneously on all branches of a bare plant or tree. Young leaves often have less [[chlorophyll]] and the leaf flush may be white or red, the latter due to presence of pigments, particularly [[Anthocyanin|anthocyanins]].<ref name=":0">{{Cite journal|last=Dominy|first=Nathaniel J.|last2=Lucas|first2=Peter W.|last3=Ramsden|first3=Lawrence W.|last4=Riba-Hernandez|first4=Pablo|last5=Stoner|first5=Kathryn E.|last6=Turner|first6=Ian M.|date=2002|title=Why are young leaves red?|url=http://doi.wiley.com/10.1034/j.1600-0706.2002.980117.x|journal=Oikos|language=en|volume=98|issue=1|pages=163–176|doi=10.1034/j.1600-0706.2002.980117.x|via=}}</ref> Leaf flushing succeeds [[Abscission#In plants|leaf fall]], and is delayed by winter in the temperate zone or by extreme dryness in the tropics. Leaf fall and leaf flushing in tropical [[deciduous]] forests can overlap in some species, called ''leaf-exchanging'' species, producing new leaves during the same period when old leaves are shed or almost immediately after.<ref name="Elliott">{{cite journal|last1=Elliott|first1=Stephen|last2=Baker|first2=Patrick J.|last3=Borchert|first3=Rolf|title=Leaf flushing during the dry season: the paradox of Asian monsoon forests|journal=Global Ecology and Biogeography|date=May 2006|volume=15|issue=3|pages=248–257|doi=10.1111/j.1466-8238.2006.00213.x|url=http://ctfs.si.edu/Public/pdfs/ToDelete/Elliott_etal_2006_GlobalEcolBiogeogr.pdf}}</ref><ref>{{cite journal|last1=Heatwole|first1=Harold|last2=Lowman|first2=Margaret D.|last3=Donovan|first3=Christopher|last4=McCoy|first4=Maria|title=Phenology of leaf-flushing and macroarthropod abundances in canopies of ''Eucalyptus'' saplings|journal=Selbyana|date=1 January 1997|volume=18|issue=2|pages=200–214|jstor=41760435}}</ref> Leaf-flushing may be synchronized among trees of a single species or even across species in an area.<ref>{{cite journal|doi=10.2307/2388052 |jstor=2388052 |title=The Causes and Consequences of Synchronous Flushing in a Dry Tropical Forest |journal=Biotropica |volume=16 |issue=3 |pages=193 |year=1984 |last1=Lieberman |first1=Diana |last2=Lieberman |first2=Milton }}</ref> In the seasonal tropics, leaf flushing [[phenology]] may be influenced by herbivory and water stress.<ref>{{cite journal|last1=Murali|first1=K. S.|last2=Sukumar|first2=R.|title=Leaf flushing phenology and herbivory in a tropical dry deciduous forest, southern India|journal=Oecologia|date=1993|volume=94|issue=1|pages=114–119|url=http://www.geocities.ws/murali_kallur/oecologia.pdf|doi=10.1007/BF00317311|pmid=28313868}}</ref><ref>{{Cite journal|last=Elliott|first=Stephen|last2=Baker|first2=Patrick J.|last3=Borchert|first3=Rolf|date=2006|title=Leaf Flushing during the Dry Season: The Paradox of Asian Monsoon Forests|url=https://www.jstor.org/stable/3697587|journal=Global Ecology and Biogeography|volume=15|issue=3|pages=248–257|issn=1466-822X}}</ref>

== Red leaf flush ==
In tropical regions, leaves often flush red when young and in the phase of expansion to mature size. Red flushing is frequent among woody species, reported from 20-40% of the woody species in a site in Costa Rica,<ref>{{Cite journal|last=Opler|first=Paul A.|last2=Frankie|first2=Gordon W.|last3=Baker|first3=Herbert G.|date=1980|title=Comparative Phenological Studies of Treelet and Shrub Species in Tropical Wet and Dry Forests in the Lowlands of Costa Rica|url=https://www.jstor.org/stable/2259250?origin=crossref|journal=The Journal of Ecology|volume=68|issue=1|pages=167|doi=10.2307/2259250|via=}}</ref> in 36% of species in Barro Colorado Island, Panama,<ref>{{Cite journal|last=Kursar|first=Thomas A.|last2=Coley|first2=Phyllis D.|date=1992|title=Delayed Greening in Tropical Leaves: An Antiherbivore Defense?|url=https://www.jstor.org/stable/2388520?origin=crossref|journal=Biotropica|volume=24|issue=2|pages=256|doi=10.2307/2388520|via=}}</ref> about 49% of species in Kibale National Park, Uganda,<ref>{{Cite journal|last=Chapman|first=C. A.|last2=Wrangham|first2=R. W.|last3=Chapman|first3=L. J.|last4=Kennard|first4=D. K.|last5=Zanne|first5=A. E.|date=1999|title=Fruit and flower phenology at two sites in Kibale National Park, Uganda|url=https://www.cambridge.org/core/product/identifier/S0266467499000759/type/journal_article|journal=Journal of Tropical Ecology|language=en|volume=15|issue=2|pages=189–211|doi=10.1017/S0266467499000759|issn=0266-4674|via=}}</ref> and in 83 of 250 species in Southern Yunnan, China.<ref name=":1">{{Cite journal|last=Gong|first=Wei-Chang|last2=Liu|first2=Yan-Hong|last3=Wang|first3=Chuan-Ming|last4=Chen|first4=Ya-Qing|last5=Martin|first5=Konrad|last6=Meng|first6=Ling-Zeng|date=2020|title=Why Are There so Many Plant Species That Transiently Flush Young Leaves Red in the Tropics?|url=https://www.frontiersin.org/articles/10.3389/fpls.2020.00083/full|journal=Frontiers in Plant Science|language=English|volume=11|doi=10.3389/fpls.2020.00083|issn=1664-462X|pmc=PMC7041174|pmid=32133020}}</ref> The red coloration is primarily due to the presence of [[Anthocyanin|anthocyanins]]. Various hypotheses have been advanced to explain red flushing. The herbivore defense hypothesis suggests that the red coloration may make the leaves less likely to be attacked by insects as they are [[Crypsis|cryptic]] to herbivores that are blind to the red part of the spectrum.<ref name=":0" /> It has also been hypothesised that the anthocyanins may reduce light stress or fungal attacks on leaves<ref name=":0" /><ref name=":1" /><ref>{{Cite journal|last=Coley|first=Phyllis D.|last2=Aide|first2=T. Mitchell|date=1989|title=Red Coloration of Tropical Young Leaves: A Possible Antifungal Defence?|url=https://www.jstor.org/stable/2559430|journal=Journal of Tropical Ecology|volume=5|issue=3|pages=293–300|issn=0266-4674}}</ref>. A recent study in tropical forest region of China provides support for the herbivore defense hypothesis, indicating that the red coloration of young leaves protects them from attacks of herbivorous insects through chemical defense as the red leaves have high concentrations of [[Tannin|tannins]] and anthocyanins.<ref name=":1" />


==References==
==References==

Revision as of 13:24, 26 January 2021

Leaf flush in an Actinodaphne tree showing young whitish leaves turning pale green against the darker green mature leaves.
A Canarium strictum tree in red leaf flush.

Leaf flushing is the production of a flush of new leaves typically produced simultaneously on all branches of a bare plant or tree. Young leaves often have less chlorophyll and the leaf flush may be white or red, the latter due to presence of pigments, particularly anthocyanins.[1] Leaf flushing succeeds leaf fall, and is delayed by winter in the temperate zone or by extreme dryness in the tropics. Leaf fall and leaf flushing in tropical deciduous forests can overlap in some species, called leaf-exchanging species, producing new leaves during the same period when old leaves are shed or almost immediately after.[2][3] Leaf-flushing may be synchronized among trees of a single species or even across species in an area.[4] In the seasonal tropics, leaf flushing phenology may be influenced by herbivory and water stress.[5][6]

Red leaf flush

In tropical regions, leaves often flush red when young and in the phase of expansion to mature size. Red flushing is frequent among woody species, reported from 20-40% of the woody species in a site in Costa Rica,[7] in 36% of species in Barro Colorado Island, Panama,[8] about 49% of species in Kibale National Park, Uganda,[9] and in 83 of 250 species in Southern Yunnan, China.[10] The red coloration is primarily due to the presence of anthocyanins. Various hypotheses have been advanced to explain red flushing. The herbivore defense hypothesis suggests that the red coloration may make the leaves less likely to be attacked by insects as they are cryptic to herbivores that are blind to the red part of the spectrum.[1] It has also been hypothesised that the anthocyanins may reduce light stress or fungal attacks on leaves[1][10][11]. A recent study in tropical forest region of China provides support for the herbivore defense hypothesis, indicating that the red coloration of young leaves protects them from attacks of herbivorous insects through chemical defense as the red leaves have high concentrations of tannins and anthocyanins.[10]

References

  1. ^ a b c Dominy, Nathaniel J.; Lucas, Peter W.; Ramsden, Lawrence W.; Riba-Hernandez, Pablo; Stoner, Kathryn E.; Turner, Ian M. (2002). "Why are young leaves red?". Oikos. 98 (1): 163–176. doi:10.1034/j.1600-0706.2002.980117.x.
  2. ^ Elliott, Stephen; Baker, Patrick J.; Borchert, Rolf (May 2006). "Leaf flushing during the dry season: the paradox of Asian monsoon forests" (PDF). Global Ecology and Biogeography. 15 (3): 248–257. doi:10.1111/j.1466-8238.2006.00213.x.
  3. ^ Heatwole, Harold; Lowman, Margaret D.; Donovan, Christopher; McCoy, Maria (1 January 1997). "Phenology of leaf-flushing and macroarthropod abundances in canopies of Eucalyptus saplings". Selbyana. 18 (2): 200–214. JSTOR 41760435.
  4. ^ Lieberman, Diana; Lieberman, Milton (1984). "The Causes and Consequences of Synchronous Flushing in a Dry Tropical Forest". Biotropica. 16 (3): 193. doi:10.2307/2388052. JSTOR 2388052.
  5. ^ Murali, K. S.; Sukumar, R. (1993). "Leaf flushing phenology and herbivory in a tropical dry deciduous forest, southern India" (PDF). Oecologia. 94 (1): 114–119. doi:10.1007/BF00317311. PMID 28313868.
  6. ^ Elliott, Stephen; Baker, Patrick J.; Borchert, Rolf (2006). "Leaf Flushing during the Dry Season: The Paradox of Asian Monsoon Forests". Global Ecology and Biogeography. 15 (3): 248–257. ISSN 1466-822X.
  7. ^ Opler, Paul A.; Frankie, Gordon W.; Baker, Herbert G. (1980). "Comparative Phenological Studies of Treelet and Shrub Species in Tropical Wet and Dry Forests in the Lowlands of Costa Rica". The Journal of Ecology. 68 (1): 167. doi:10.2307/2259250.
  8. ^ Kursar, Thomas A.; Coley, Phyllis D. (1992). "Delayed Greening in Tropical Leaves: An Antiherbivore Defense?". Biotropica. 24 (2): 256. doi:10.2307/2388520.
  9. ^ Chapman, C. A.; Wrangham, R. W.; Chapman, L. J.; Kennard, D. K.; Zanne, A. E. (1999). "Fruit and flower phenology at two sites in Kibale National Park, Uganda". Journal of Tropical Ecology. 15 (2): 189–211. doi:10.1017/S0266467499000759. ISSN 0266-4674.
  10. ^ a b c Gong, Wei-Chang; Liu, Yan-Hong; Wang, Chuan-Ming; Chen, Ya-Qing; Martin, Konrad; Meng, Ling-Zeng (2020). "Why Are There so Many Plant Species That Transiently Flush Young Leaves Red in the Tropics?". Frontiers in Plant Science. 11. doi:10.3389/fpls.2020.00083. ISSN 1664-462X. PMC 7041174. PMID 32133020.{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)
  11. ^ Coley, Phyllis D.; Aide, T. Mitchell (1989). "Red Coloration of Tropical Young Leaves: A Possible Antifungal Defence?". Journal of Tropical Ecology. 5 (3): 293–300. ISSN 0266-4674.