Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in

plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclu ...

s and

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

s. 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.

Plants

Many

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s (angiosperms) use a

photoreceptor protein Photoreceptor proteins are light-sensitive proteins involved in the sensing and response to light in a variety of organisms. Some examples are rhodopsin in the photoreceptor cells of the vertebrate retina, phytochrome in plants, and bacterior ...

, such as

phytochrome Phytochromes are a class of photoreceptor in plants, bacteria and fungi used to detect light. They are sensitive to light in the red and far-red region of the visible spectrum and can be classed as either Type I, which are activated by far-re ...

cryptochrome Cryptochromes (from the Greek κρυπτός χρώμα, "hidden colour") are a class of flavoproteins found in plants and animals that are sensitive to blue light. They are involved in the circadian rhythms and the sensing of magnetic fields ...

, 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: P_r and P_fr. Red light (which is present during the day) converts phytochrome to its active form (pfr). This then triggers the plant to grow. In turn, far-red light is present in the shade or in the dark and this converts phytochrome from pfr to pr. P_r is the inactive form of phytochrome and will not allow for plant growth. This system of P_fr to P_r conversion allows the plant to sense when it is night and when it is day. P_fr can also be converted back to P_r by a process known as dark reversion, where long periods of darkness trigger the conversion of P_fr. 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. 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. 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. 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. 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, but it was later discovered that the length of the night was the controlling factor. 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. Each plant has a different length critical photoperiod, or critical night length. Modern biologists believe 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.

Long-day plants

Long-day plants flower when the night length falls below their critical photoperiod. 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. After that date, days grow shorter (i.e. nights grow longer) until 21 December (the

winter solstice The winter solstice, also called the hibernal solstice, occurs when either of Earth's poles reaches its maximum tilt away from the Sun. This happens twice yearly, once in each hemisphere (Northern and Southern). For that hemisphere, the winter ...

). This situation is reversed in the southern hemisphere (i.e., longest day is 21 December and shortest day is 21 June). Some long-day obligate plants are: *

Carnation ''Dianthus caryophyllus'' (), commonly known as the carnation or clove pink, is a species of ''Dianthus''. It is likely native to the Mediterranean region but its exact range is unknown due to extensive cultivation for the last 2,000 years.Med ...

(''Dianthus'') *

Henbane ''Hyoscyamus niger'', commonly known as henbane, black henbane, or stinking nightshade, is a poisonous plant in the nightshade family Solanaceae. It is native to temperate Europe and Siberia, and naturalised in Great Britain and Ireland. ...

(''Hyoscyamus'') *

Oat The oat (''Avena sativa''), sometimes called the common oat, is a species of cereal grain grown for its seed, which is known by the same name (usually in the plural, unlike other cereals and pseudocereals). While oats are suitable for human con ...

(''Avena'') Some long-day facultative plants are: *

Pea The pea is most commonly the small spherical seed or the seed-pod of the flowering plant species ''Pisum sativum''. Each pod contains several peas, which can be green or yellow. Botanically, pea pods are fruit, since they contain seeds and d ...

(''Pisum sativum) *

Barley Barley (''Hordeum vulgare''), a member of the grass family, is a major cereal grain grown in temperate climates globally. It was one of the first cultivated grains, particularly in Eurasia as early as 10,000 years ago. Globally 70% of barley p ...

(''Hordeum vulgare'') *

Lettuce Lettuce (''Lactuca sativa'') is an annual plant of the family Asteraceae. It is most often grown as a leaf vegetable, but sometimes for its stem and seeds. Lettuce is most often used for salads, although it is also seen in other kinds of food, ...

(''Lactuca sativa'') *

Wheat Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeologi ...

(''Triticum aestivum'')

Short-day plants

Short-day plants flower when the night lengths exceed their critical photoperiod. 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. In general, short-day (i.e. long-night) 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: *

Kenaf Kenaf tymology: Persian ''Hibiscus cannabinus'', is a plant in the family Malvaceae also called Deccan hemp and Java jute. ''Hibiscus cannabinus'' is in the genus '' Hibiscus'' and is native to Africa, though its exact origin is unknown. The name ...

( ''Hibiscus cannabinus'') * Marijuana (''Cannabis'') *

Cotton Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus '' Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor pe ...

(''Gossypium'') *

Rice Rice is the seed of the grass species '' Oryza sativa'' (Asian rice) or less commonly ''Oryza glaberrima'' (African rice). The name wild rice is usually used for species of the genera '' Zizania'' and '' Porteresia'', both wild and domesticat ...

(''Oryza'') * Sorghum (''Sorghum bicolor'') * Green gram (Mung bean, ''Vigna radiata'') *

Soybean The soybean, soy bean, or soya bean (''Glycine max'') is a species of legume native to East Asia, widely grown for its edible bean, which has numerous uses. Traditional unfermented food uses of soybeans include soy milk, from which tofu a ...

s (''Glycine max'')

Day-neutral plants

Day-neutral plants, such as cucumbers,

rose A rose is either a woody perennial flowering plant of the genus ''Rosa'' (), in the family Rosaceae (), or the flower it bears. There are over three hundred species and tens of thousands of cultivars. They form a group of plants that can be ...

tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...

es, and Ruderalis (

autoflowering cannabis Autoflowering cannabis or day neutral cannabis varieties automatically switch from vegetative growth to the flowering stage based on age, as opposed to the ratio of light to dark hours required with photoperiod dependent/short-day strains. Ma ...

) do not initiate flowering based on photoperiodism. 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).

Animals

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 Human sexual activity, human sexual practice or human sexual behaviour is the manner in which humans experience and express their sexuality. People engage in a variety of sexual acts, ranging from activities done alone (e.g., masturbation) ...

, and even the resizing of sexual organs. The singing frequency of birds such as the

canary Canary originally referred to the island of Gran Canaria on the west coast of Africa, and the group of surrounding islands (the Canary Islands). It may also refer to: Animals Birds * Canaries, birds in the genera ''Serinus'' and ''Crithagra'' i ...

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 (formerly, hyperstriatum ventrale, pars caudalis (HVc), and high vocal center) is a nucleus in the brain of the songbirds (order passeriformes) necessary for both the learning and the production of bird song. It is located in the lateral cauda ...

(HVC) and the robust nucleus of the archistriatum (RA) increase in size. When the photoperiod decreases, these areas of the brain regress. 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 In neuroanatomy, the retinohypothalamic tract (RHT) is a photic neural input pathway involved in the circadian rhythms of mammals. The origin of the retinohypothalamic tract is the intrinsically photosensitive retinal ganglion cells (ipRGC), wh ...

(RHT). Some mammals are highly seasonal, while humans' seasonality is largely believed to be evolutionary baggage.

Plants

Long-day plants

Short-day plants

Day-neutral plants

Animals

See also

References

Further reading