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Photoperiodism

February 15, 2024

Photoperiodism is the physiological reaction of organisms to the length of light or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light and dark periods. They are classified under three groups according to the photoperiods: short-day plants, long-day plants, and day-neutral plants.

In animals photoperiodism (sometimes called seasonality) is the suite of physiological changes that occur in response to changes in day length. This allows animals to respond to a temporally changing environment associated with changing seasons as the earth orbits the sun.

Plants edit

 
Pr converts to Pfr during the day time and Pfr slowly reverts to Pr during the night time. When nights are short, an excess amount of Pfr remains in the day time and during long nights, most of the Pfr is reverted to Pr.

Many flowering plants (angiosperms) use a circadian rhythm together with photoreceptor protein, such as phytochrome or cryptochrome,[1] to sense seasonal changes in night length, or photoperiod, which they take as signals to flower. In a further subdivision, obligate photoperiodic plants absolutely require a long or short enough night before flowering, whereas facultative photoperiodic plants are more likely to flower under one condition.

Phytochrome comes in two forms: Pr and Pfr. Red light (which is present during the day) converts phytochrome to its active form (Pfr) which then stimulates various processes such as germination, flowering or branching. In comparison, plants receive more far-red in the shade, and this converts phytochrome from Pfr to its inactive form, Pr, inhibiting germination. This system of Pfr to Pr conversion allows the plant to sense when it is night and when it is day.[2] Pfr can also be converted back to Pr by a process known as dark reversion, where long periods of darkness trigger the conversion of Pfr.[3] This is important in regards to plant flowering. Experiments by Halliday et al. showed that manipulations of the red-to far-red ratio in Arabidopsis can alter flowering. They discovered that plants tend to flower later when exposed to more red light, proving that red light is inhibitory to flowering.[4] Other experiments have proven this by exposing plants to extra red-light in the middle of the night. A short-day plant will not flower if light is turned on for a few minutes in the middle of the night and a long-day plant can flower if exposed to more red-light in the middle of the night.[5]

Cryptochromes are another type of photoreceptor that is important in photoperiodism. Cryptochromes absorb blue light and UV-A. Cryptochromes entrain the circadian clock to light.[6] It has been found that both cryptochrome and phytochrome abundance relies on light and the amount of cryptochrome can change depending on day-length. This shows how important both of the photoreceptors are in regards to determining day-length.[7]

In 1920, W. W. Garner and H. A. Allard published their discoveries on photoperiodism and felt it was the length of daylight that was critical,[1][8] but it was later discovered that the length of the night was the controlling factor.[9][10] Photoperiodic flowering plants are classified as long-day plants or short-day plants even though night is the critical factor because of the initial misunderstanding about daylight being the controlling factor. Along with long-day plants and short-day plants, there are plants that fall into a "dual-day length category". These plants are either long-short-day plants (LSDP) or short-long-day plants (SLDP). LSDPs flower after a series of long days followed by short days whereas SLDPs flower after a series of short days followed by long days.[11] Each plant has a different length critical photoperiod, or critical night length.[1]

Modern biologists believe[12] that it is the coincidence of the active forms of phytochrome or cryptochrome, created by light during the daytime, with the rhythms of the circadian clock that allows plants to measure the length of the night. Other than flowering, photoperiodism in plants includes the growth of stems or roots during certain seasons and the loss of leaves. Artificial lighting can be used to induce extra-long days.[1]

Long-day plants edit

 

Long-day plants flower when the night length falls below their critical photoperiod.[13] These plants typically flower during late spring or early summer as days are getting longer. In the northern hemisphere, the longest day of the year (summer solstice) is on or about 21 June.[14] After that date, days grow shorter (i.e. nights grow longer) until 21 December (the winter solstice). This situation is reversed in the southern hemisphere (i.e., longest day is 21 December and shortest day is 21 June).[1][8]

Some long-day obligate plants are:

Some long-day facultative plants are:

Short-day plants edit

 

Short-day (also called long-night) plants flower when the night lengths exceed their critical photoperiod.[15] They cannot flower under short nights or if a pulse of artificial light is shone on the plant for several minutes during the night; they require a continuous period of darkness before floral development can begin. Natural nighttime light, such as moonlight or lightning, is not of sufficient brightness or duration to interrupt flowering.[1][8]

Short-day plants flower as days grow shorter (and nights grow longer) after 21 June in the northern hemisphere, which is during summer or fall. The length of the dark period required to induce flowering differs among species and varieties of a species.

Photoperiodism affects flowering by inducing the shoot to produce floral buds instead of leaves and lateral buds.

Some short-day facultative plants are:[16]

Day-neutral plants edit

Day-neutral plants, such as cucumbers, roses, tomatoes, and Ruderalis (autoflowering cannabis) do not initiate flowering based on photoperiodism.[18] Instead, they may initiate flowering after attaining a certain overall developmental stage or age, or in response to alternative environmental stimuli, such as vernalisation (a period of low temperature).[1][8]

Animals edit

Daylength, and thus knowledge of the season of the year, is vital to many animals. A number of biological and behavioural changes are dependent on this knowledge. Together with temperature changes, photoperiod provokes changes in the color of fur and feathers, migration, entry into hibernation, sexual behaviour, and even the resizing of organs.

In insects, sensitivity to photoperiod has been proven to be initiated by photoreceptors located in the brain.[19][20] Photoperiod can affect insects at different life stages, serving as an environmental cue for physiological processes such as diapause induction and termination, and seasonal morphs.[21] In the water strider Aquarius paludum, for instance, photoperiod conditions during nymphal development have been shown to trigger seasonal changes in wing frequency and also induce diapause, although the threshold critical day lengths for the determination of both traits diverged by about an hour.[22] In Gerris buenoi, another water strider species, photoperiod has also been shown to be the cause of wing polyphenism,[23] although the specific daylengths changed between species, suggesting that phenotypic plasticity in response to photoperiod has evolved even between relatively closely related species.

The singing frequency of birds such as the canary depends on the photoperiod. In the spring, when the photoperiod increases (more daylight), the male canary's testes grow. As the testes grow, more androgens are secreted and song frequency increases. During autumn, when the photoperiod decreases (less daylight), the male canary's testes regress and androgen levels drop dramatically, resulting in decreased singing frequency. Not only is singing frequency dependent on the photoperiod but the song repertoire is also. The long photoperiod of spring results in a greater song repertoire. Autumn's shorter photoperiod results in a reduction in song repertoire. These behavioral photoperiod changes in male canaries are caused by changes in the song center of the brain. As the photoperiod increases, the high vocal center (HVC) and the robust nucleus of the archistriatum (RA) increase in size. When the photoperiod decreases, these areas of the brain regress.[24]

In mammals, daylength is registered in the suprachiasmatic nucleus (SCN), which is informed by retinal light-sensitive ganglion cells, which are not involved in vision. The information travels through the retinohypothalamic tract (RHT). In most species the hormone melatonin is produced by the pineal gland only during the hours of darkness, influenced by the light input through the RHT and by innate circadian rhythms. This hormonal signal, combined with outputs from the SCN inform the rest of the body about the time of day, and the length of time that melatonin is secreted is how the time of year is perceived.

Some mammals are highly seasonal. The view has been expressed that humans' seasonality is largely believed to be evolutionary baggage.[25][relevant?]. Human birth rate varies throughout the year, and the peak month of births appears to vary by latitude.[26] Seasonality in human birth rate appears to have largely decreased since the industrial revolution.[27][28]

Other organisms edit

Photoperiodism has also been demonstrated in other organisms besides plants and animals. The fungus Neurospora crassa as well as the dinoflagellate Lingulodinium polyedra and the unicellular green alga Chlamydomonas reinhardtii have been shown to display photoperiodic responses[29][30][31].

See also edit

References edit

  1. ^ a b c d e f g Mauseth JD (2003). Botany : An Introduction to Plant Biology (3rd ed.). Sudbury, MA: Jones and Bartlett Learning. pp. 422–27. ISBN 978-0-7637-2134-3.
  2. ^ Fankhauser C (April 2001). "The phytochromes, a family of red/far-red absorbing photoreceptors". The Journal of Biological Chemistry. 276 (15): 11453–11456. doi:10.1074/jbc.R100006200. PMID 11279228.
  3. ^ Casal JJ, Candia AN, Sellaro R (June 2014). "Light perception and signalling by phytochrome A". Journal of Experimental Botany. 65 (11): 2835–2845. doi:10.1093/jxb/ert379. hdl:11336/4338. PMID 24220656.
  4. ^ Lin C (May 2000). "Photoreceptors and regulation of flowering time". Plant Physiology. 123 (1): 39–50. doi:10.1104/pp.123.1.39. PMC 1539253. PMID 10806223.
  5. ^ Chamovitz D (2013). What A Plant Knows. Scientific American. pp. 17–18. ISBN 978-0-374-28873-0.
  6. ^ Lin C, Todo T (2005). "The cryptochromes". Genome Biology. 6 (5): 220. doi:10.1186/gb-2005-6-5-220. PMC 1175950. PMID 15892880.
  7. ^ Mockler T, Yang H, Yu X, Parikh D, Cheng YC, Dolan S, Lin C (February 2003). "Regulation of photoperiodic flowering by Arabidopsis photoreceptors". Proceedings of the National Academy of Sciences of the United States of America. 100 (4): 2140–2145. Bibcode:2003PNAS..100.2140M. doi:10.1073/pnas.0437826100. PMC 149972. PMID 12578985.
  8. ^ a b c d Capon B (2005). Botany for Gardeners (2nd ed.). Portland, OR: Timber Publishing. pp. 148–51. ISBN 978-0-88192-655-2.
  9. ^ Hamner KC, Bonner J (1938). "Photoperiodism in relation to hormones as factors in floral initiation and development" (PDF). Botanical Gazette. 100 (2): 388–431. doi:10.1086/334793. JSTOR 2471641. S2CID 84084837.
  10. ^ Hamner KC (1940). "Interrelation of light and darkness in photoperiodic induction". Botanical Gazette. 101 (3): 658–87. doi:10.1086/334903. JSTOR 2472399. S2CID 83682483.
  11. ^ Taiz L, Zeiger E, Møller I, Murphy A (2015). Plant Physiology and Development (Sixth ed.). Sunderland, MA: Sinauer Associates, Inc. ISBN 978-1-60535-353-1.
  12. ^ Andrés F, Galbraith DW, Talón M, Domingo C (October 2009). "Analysis of PHOTOPERIOD SENSITIVITY5 sheds light on the role of phytochromes in photoperiodic flowering in rice". Plant Physiology. 151 (2): 681–690. doi:10.1104/pp.109.139097. PMC 2754645. PMID 19675157.
  13. ^ Starr C, Taggart R, Evers C, Starr L (2013). Plant Structure and Function. Vol. 4 (13th ed.). Brooks/Cole. p. 517. ISBN 978-1-111-58068-1.
  14. ^ Gooley T (2010-03-30). The Natural Navigator. Random House. ISBN 978-0-7535-2311-7.
  15. ^ BSCS Biology (9 ed.). BSCS. 2002. p. 519. ISBN 978-0-7872-9008-5.
  16. ^ Jones HG (1992). Plants and Microclimate: A Quantitative Approach to Environmental Plant Physiology. Cambridge University Press. p. 225. ISBN 978-0-521-42524-7.
  17. ^ Purcell LC, Salmeron M, Ashlock L (2014). "Chapter 2" (PDF). Arkansas Soybean Production Handbook - MP197. Little Rock, Arkansas: University of Arkansas Cooperative Extension Service. pp. 5–7. Retrieved 21 February 2016.
  18. ^ Meneely P (2014). Genetic Analysis: Genes, Genomes, and Networks in Eukaryotes (2nd ed.). Oxford University Press. p. 373. ISBN 978-0-19-968126-6.
  19. ^ Claret J (1966). "Recherche du centre photorecepteur lors de l'induction de la diapause chez Pieris brassicae L.". Comptes Rendus de l'Académie des Sciences. 262: 553–556.
  20. ^ Bowen MF, Saunders DS, Bollenbacher WE, Gilbert LI (September 1984). "In vitro reprogramming of the photoperiodic clock in an insect brain-retrocerebral complex". Proceedings of the National Academy of Sciences of the United States of America. 81 (18): 5881–4. Bibcode:1984PNAS...81.5881B. doi:10.1073/pnas.81.18.5881. PMC 391816. PMID 6592591.
  21. ^ Saunders DS (2012). "Insect photoperiodism: seeing the light". Physiological Entomology. 37 (3): 207–218. doi:10.1111/j.1365-3032.2012.00837.x. S2CID 85249708.
  22. ^ Harada T, Numata H (1993). "Two Critical Day Lengths for the Determination of Wing Forms and the Induction of Adult Diapause in the Water Strider, Aquarius paludum". Naturwissenschaften. 80 (9): 430–432. Bibcode:1993NW.....80..430H. doi:10.1007/BF01168342. S2CID 39616943.
  23. ^ Gudmunds E, Narayanan S, Lachivier E, Duchemin M, Khila A, Husby A (April 2022). "Photoperiod controls wing polyphenism in a water strider independently of insulin receptor signalling". Proceedings. Biological Sciences. 289 (1973): 20212764. doi:10.1098/rspb.2021.2764. PMC 9043737. PMID 35473377.
  24. ^ Nelson RJ (2005). An Introduction to Behavioral Endocrinology. Sunderland, MA: Sinauer Associates. p. 189.
  25. ^ Foster R, Williams R (5 December 2009). "Extra-retinal photo receptors" (Interview). Science Show. ABC Radio National. Retrieved 2010-05-28. ...we have the evolutionary baggage of showing seasonality but we're not entirely sure what the mechanism is.
  26. ^ Martinez-Bakker, Micaela; Bakker, Kevin M.; King, Aaron A.; Rohani, Pejman (2014-05-22). "Human birth seasonality: latitudinal gradient and interplay with childhood disease dynamics". Proceedings of the Royal Society B: Biological Sciences. 281 (1783): 20132438. doi:10.1098/rspb.2013.2438. ISSN 0962-8452. PMC 3996592. PMID 24695423.
  27. ^ Martinez-Bakker, Micaela; Bakker, Kevin M.; King, Aaron A.; Rohani, Pejman (2014-05-22). "Human birth seasonality: latitudinal gradient and interplay with childhood disease dynamics". Proceedings of the Royal Society B: Biological Sciences. 281 (1783): 20132438. doi:10.1098/rspb.2013.2438. ISSN 0962-8452. PMC 3996592. PMID 24695423.
  28. ^ Wehr, Thomas A. (August 2001). "Photoperiodism in Humans and Other Primates: Evidence and Implications". Journal of Biological Rhythms. 16 (4): 348–364. doi:10.1177/074873001129002060. ISSN 0748-7304. PMID 11506380. S2CID 25886221.
  29. ^ Tan Y, Merrow M, Roenneberg T. Photoperiodism in Neurospora crassa. J Biol Rhythms. 2004 Apr;19(2):135-43. https://doi.org/10.1177/0748730404263015. PMID: 15038853.
  30. ^ Suzuki, L., Johnson, C. Photoperiodic control of germination in the unicell Chlamydomonas. Naturwissenschaften 89, 214–220 (2002). https://doi.org/10.1007/s00114-002-0302-6
  31. ^ Ivonne Balzer, Rüdiger Hardeland ,Photoperiodism and Effects of Indoleamines in a Unicellular Alga, Gonyaulax polyedra.Science253,795-797(1991).https://doi.org/10.1126/science.1876838

Further reading edit

  • Fosket DE (1994). Plant Growth & Development, A Molecular Approach. San Diego: Academic Press. p. 495.
  • Thomas B, Vince-Prue D (1997). Photoperiodism in plants (2nd ed.). Academic Press.

February 15, 2024
photoperiodism, this, article, lead, section, short, adequately, summarize, points, please, consider, expanding, lead, provide, accessible, overview, important, aspects, article, october, 2023, physiological, reaction, organisms, length, light, dark, period, o. This article s lead section may be too short to adequately summarize the key points Please consider expanding the lead to provide an accessible overview of all important aspects of the article October 2023 Photoperiodism is the physiological reaction of organisms to the length of light or a dark period It occurs in plants and animals Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light and dark periods They are classified under three groups according to the photoperiods short day plants long day plants and day neutral plants In animals photoperiodism sometimes called seasonality is the suite of physiological changes that occur in response to changes in day length This allows animals to respond to a temporally changing environment associated with changing seasons as the earth orbits the sun Contents 1 Plants 1 1 Long day plants 1 2 Short day plants 1 3 Day neutral plants 2 Animals 3 Other organisms 4 See also 5 References 6 Further readingPlants edit nbsp Pr converts to Pfr during the day time and Pfr slowly reverts to Pr during the night time When nights are short an excess amount of Pfr remains in the day time and during long nights most of the Pfr is reverted to Pr Many flowering plants angiosperms use a circadian rhythm together with photoreceptor protein such as phytochrome or cryptochrome 1 to sense seasonal changes in night length or photoperiod which they take as signals to flower In a further subdivision obligate photoperiodic plants absolutely require a long or short enough night before flowering whereas facultative photoperiodic plants are more likely to flower under one condition Phytochrome comes in two forms Pr and Pfr Red light which is present during the day converts phytochrome to its active form Pfr which then stimulates various processes such as germination flowering or branching In comparison plants receive more far red in the shade and this converts phytochrome from Pfr to its inactive form Pr inhibiting germination This system of Pfr to Pr conversion allows the plant to sense when it is night and when it is day 2 Pfr can also be converted back to Pr by a process known as dark reversion where long periods of darkness trigger the conversion of Pfr 3 This is important in regards to plant flowering Experiments by Halliday et al showed that manipulations of the red to far red ratio in Arabidopsis can alter flowering They discovered that plants tend to flower later when exposed to more red light proving that red light is inhibitory to flowering 4 Other experiments have proven this by exposing plants to extra red light in the middle of the night A short day plant will not flower if light is turned on for a few minutes in the middle of the night and a long day plant can flower if exposed to more red light in the middle of the night 5 Cryptochromes are another type of photoreceptor that is important in photoperiodism Cryptochromes absorb blue light and UV A Cryptochromes entrain the circadian clock to light 6 It has been found that both cryptochrome and phytochrome abundance relies on light and the amount of cryptochrome can change depending on day length This shows how important both of the photoreceptors are in regards to determining day length 7 In 1920 W W Garner and H A Allard published their discoveries on photoperiodism and felt it was the length of daylight that was critical 1 8 but it was later discovered that the length of the night was the controlling factor 9 10 Photoperiodic flowering plants are classified as long day plants or short day plants even though night is the critical factor because of the initial misunderstanding about daylight being the controlling factor Along with long day plants and short day plants there are plants that fall into a dual day length category These plants are either long short day plants LSDP or short long day plants SLDP LSDPs flower after a series of long days followed by short days whereas SLDPs flower after a series of short days followed by long days 11 Each plant has a different length critical photoperiod or critical night length 1 Modern biologists believe 12 that it is the coincidence of the active forms of phytochrome or cryptochrome created by light during the daytime with the rhythms of the circadian clock that allows plants to measure the length of the night Other than flowering photoperiodism in plants includes the growth of stems or roots during certain seasons and the loss of leaves Artificial lighting can be used to induce extra long days 1 Long day plants edit nbsp Long day plants flower when the night length falls below their critical photoperiod 13 These plants typically flower during late spring or early summer as days are getting longer In the northern hemisphere the longest day of the year summer solstice is on or about 21 June 14 After that date days grow shorter i e nights grow longer until 21 December the winter solstice This situation is reversed in the southern hemisphere i e longest day is 21 December and shortest day is 21 June 1 8 Some long day obligate plants are Carnation Dianthus Henbane Hyoscyamus Oat Avena Some long day facultative plants are Pea Pisum sativum Barley Hordeum vulgare Lettuce Lactuca sativa Wheat Triticum aestivum Short day plants edit nbsp Short day also called long night plants flower when the night lengths exceed their critical photoperiod 15 They cannot flower under short nights or if a pulse of artificial light is shone on the plant for several minutes during the night they require a continuous period of darkness before floral development can begin Natural nighttime light such as moonlight or lightning is not of sufficient brightness or duration to interrupt flowering 1 8 Short day plants flower as days grow shorter and nights grow longer after 21 June in the northern hemisphere which is during summer or fall The length of the dark period required to induce flowering differs among species and varieties of a species Photoperiodism affects flowering by inducing the shoot to produce floral buds instead of leaves and lateral buds Some short day facultative plants are 16 Kenaf Hibiscus cannabinus Marijuana Cannabis Cotton Gossypium Rice Oryza Sorghum Sorghum bicolor Green gram Mung bean Vigna radiata Soybeans 17 Glycine max Day neutral plants edit Day neutral plants such as cucumbers roses tomatoes and Ruderalis autoflowering cannabis do not initiate flowering based on photoperiodism 18 Instead they may initiate flowering after attaining a certain overall developmental stage or age or in response to alternative environmental stimuli such as vernalisation a period of low temperature 1 8 Animals editDaylength and thus knowledge of the season of the year is vital to many animals A number of biological and behavioural changes are dependent on this knowledge Together with temperature changes photoperiod provokes changes in the color of fur and feathers migration entry into hibernation sexual behaviour and even the resizing of organs In insects sensitivity to photoperiod has been proven to be initiated by photoreceptors located in the brain 19 20 Photoperiod can affect insects at different life stages serving as an environmental cue for physiological processes such as diapause induction and termination and seasonal morphs 21 In the water strider Aquarius paludum for instance photoperiod conditions during nymphal development have been shown to trigger seasonal changes in wing frequency and also induce diapause although the threshold critical day lengths for the determination of both traits diverged by about an hour 22 In Gerris buenoi another water strider species photoperiod has also been shown to be the cause of wing polyphenism 23 although the specific daylengths changed between species suggesting that phenotypic plasticity in response to photoperiod has evolved even between relatively closely related species The singing frequency of birds such as the canary depends on the photoperiod In the spring when the photoperiod increases more daylight the male canary s testes grow As the testes grow more androgens are secreted and song frequency increases During autumn when the photoperiod decreases less daylight the male canary s testes regress and androgen levels drop dramatically resulting in decreased singing frequency Not only is singing frequency dependent on the photoperiod but the song repertoire is also The long photoperiod of spring results in a greater song repertoire Autumn s shorter photoperiod results in a reduction in song repertoire These behavioral photoperiod changes in male canaries are caused by changes in the song center of the brain As the photoperiod increases the high vocal center HVC and the robust nucleus of the archistriatum RA increase in size When the photoperiod decreases these areas of the brain regress 24 In mammals daylength is registered in the suprachiasmatic nucleus SCN which is informed by retinal light sensitive ganglion cells which are not involved in vision The information travels through the retinohypothalamic tract RHT In most species the hormone melatonin is produced by the pineal gland only during the hours of darkness influenced by the light input through the RHT and by innate circadian rhythms This hormonal signal combined with outputs from the SCN inform the rest of the body about the time of day and the length of time that melatonin is secreted is how the time of year is perceived Some mammals are highly seasonal The view has been expressed that humans seasonality is largely believed to be evolutionary baggage 25 relevant Human birth rate varies throughout the year and the peak month of births appears to vary by latitude 26 Seasonality in human birth rate appears to have largely decreased since the industrial revolution 27 28 Other organisms editPhotoperiodism has also been demonstrated in other organisms besides plants and animals The fungus Neurospora crassa as well as the dinoflagellate Lingulodinium polyedra and the unicellular green alga Chlamydomonas reinhardtii have been shown to display photoperiodic responses 29 30 31 See also editChronobiology Circadian clock Circadian rhythm Florigen Photobiology Seasonal Breeder ScotobiologyReferences edit a b c d e f g Mauseth JD 2003 Botany An Introduction to Plant Biology 3rd ed Sudbury MA Jones and Bartlett Learning pp 422 27 ISBN 978 0 7637 2134 3 Fankhauser C April 2001 The phytochromes a family of red far red absorbing photoreceptors The Journal of Biological Chemistry 276 15 11453 11456 doi 10 1074 jbc R100006200 PMID 11279228 Casal JJ Candia AN Sellaro R June 2014 Light perception and signalling by phytochrome A Journal of Experimental Botany 65 11 2835 2845 doi 10 1093 jxb ert379 hdl 11336 4338 PMID 24220656 Lin C May 2000 Photoreceptors and regulation of flowering time Plant Physiology 123 1 39 50 doi 10 1104 pp 123 1 39 PMC 1539253 PMID 10806223 Chamovitz D 2013 What A Plant Knows Scientific American pp 17 18 ISBN 978 0 374 28873 0 Lin C Todo T 2005 The cryptochromes Genome Biology 6 5 220 doi 10 1186 gb 2005 6 5 220 PMC 1175950 PMID 15892880 Mockler T Yang H Yu X Parikh D Cheng YC Dolan S Lin C February 2003 Regulation of photoperiodic flowering by Arabidopsis photoreceptors Proceedings of the National Academy of Sciences of the United States of America 100 4 2140 2145 Bibcode 2003PNAS 100 2140M doi 10 1073 pnas 0437826100 PMC 149972 PMID 12578985 a b c d Capon B 2005 Botany for Gardeners 2nd ed Portland OR Timber Publishing pp 148 51 ISBN 978 0 88192 655 2 Hamner KC Bonner J 1938 Photoperiodism in relation to hormones as factors in floral initiation and development PDF Botanical Gazette 100 2 388 431 doi 10 1086 334793 JSTOR 2471641 S2CID 84084837 Hamner KC 1940 Interrelation of light and darkness in photoperiodic induction Botanical Gazette 101 3 658 87 doi 10 1086 334903 JSTOR 2472399 S2CID 83682483 Taiz L Zeiger E Moller I Murphy A 2015 Plant Physiology and Development Sixth ed Sunderland MA Sinauer Associates Inc ISBN 978 1 60535 353 1 Andres F Galbraith DW Talon M Domingo C October 2009 Analysis of PHOTOPERIOD SENSITIVITY5 sheds light on the role of phytochromes in photoperiodic flowering in rice Plant Physiology 151 2 681 690 doi 10 1104 pp 109 139097 PMC 2754645 PMID 19675157 Starr C Taggart R Evers C Starr L 2013 Plant Structure and Function Vol 4 13th ed Brooks Cole p 517 ISBN 978 1 111 58068 1 Gooley T 2010 03 30 The Natural Navigator Random House ISBN 978 0 7535 2311 7 BSCS Biology 9 ed BSCS 2002 p 519 ISBN 978 0 7872 9008 5 Jones HG 1992 Plants and Microclimate A Quantitative Approach to Environmental Plant Physiology Cambridge University Press p 225 ISBN 978 0 521 42524 7 Purcell LC Salmeron M Ashlock L 2014 Chapter 2 PDF Arkansas Soybean Production Handbook MP197 Little Rock Arkansas University of Arkansas Cooperative Extension Service pp 5 7 Retrieved 21 February 2016 Meneely P 2014 Genetic Analysis Genes Genomes and Networks in Eukaryotes 2nd ed Oxford University Press p 373 ISBN 978 0 19 968126 6 Claret J 1966 Recherche du centre photorecepteur lors de l induction de la diapause chez Pieris brassicae L Comptes Rendus de l Academie des Sciences 262 553 556 Bowen MF Saunders DS Bollenbacher WE Gilbert LI September 1984 In vitro reprogramming of the photoperiodic clock in an insect brain retrocerebral complex Proceedings of the National Academy of Sciences of the United States of America 81 18 5881 4 Bibcode 1984PNAS 81 5881B doi 10 1073 pnas 81 18 5881 PMC 391816 PMID 6592591 Saunders DS 2012 Insect photoperiodism seeing the light Physiological Entomology 37 3 207 218 doi 10 1111 j 1365 3032 2012 00837 x S2CID 85249708 Harada T Numata H 1993 Two Critical Day Lengths for the Determination of Wing Forms and the Induction of Adult Diapause in the Water Strider Aquarius paludum Naturwissenschaften 80 9 430 432 Bibcode 1993NW 80 430H doi 10 1007 BF01168342 S2CID 39616943 Gudmunds E Narayanan S Lachivier E Duchemin M Khila A Husby A April 2022 Photoperiod controls wing polyphenism in a water strider independently of insulin receptor signalling Proceedings Biological Sciences 289 1973 20212764 doi 10 1098 rspb 2021 2764 PMC 9043737 PMID 35473377 Nelson RJ 2005 An Introduction to Behavioral Endocrinology Sunderland MA Sinauer Associates p 189 Foster R Williams R 5 December 2009 Extra retinal photo receptors Interview Science Show ABC Radio National Retrieved 2010 05 28 we have the evolutionary baggage of showing seasonality but we re not entirely sure what the mechanism is Martinez Bakker Micaela Bakker Kevin M King Aaron A Rohani Pejman 2014 05 22 Human birth seasonality latitudinal gradient and interplay with childhood disease dynamics Proceedings of the Royal Society B Biological Sciences 281 1783 20132438 doi 10 1098 rspb 2013 2438 ISSN 0962 8452 PMC 3996592 PMID 24695423 Martinez Bakker Micaela Bakker Kevin M King Aaron A Rohani Pejman 2014 05 22 Human birth seasonality latitudinal gradient and interplay with childhood disease dynamics Proceedings of the Royal Society B Biological Sciences 281 1783 20132438 doi 10 1098 rspb 2013 2438 ISSN 0962 8452 PMC 3996592 PMID 24695423 Wehr Thomas A August 2001 Photoperiodism in Humans and Other Primates Evidence and Implications Journal of Biological Rhythms 16 4 348 364 doi 10 1177 074873001129002060 ISSN 0748 7304 PMID 11506380 S2CID 25886221 Tan Y Merrow M Roenneberg T Photoperiodism in Neurospora crassa J Biol Rhythms 2004 Apr 19 2 135 43 https doi org 10 1177 0748730404263015 PMID 15038853 Suzuki L Johnson C Photoperiodic control of germination in the unicell Chlamydomonas Naturwissenschaften 89 214 220 2002 https doi org 10 1007 s00114 002 0302 6 Ivonne Balzer Rudiger Hardeland Photoperiodism and Effects of Indoleamines in a Unicellular Alga Gonyaulax polyedra Science253 795 797 1991 https doi org 10 1126 science 1876838Further reading editFosket DE 1994 Plant Growth amp Development A Molecular Approach San Diego Academic Press p 495 Thomas B Vince Prue D 1997 Photoperiodism in plants 2nd ed Academic Press Retrieved from https en wikipedia org w index php title Photoperiodism amp oldid 1205249775, wikipedia, wiki, book, books, library,

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