Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs in plants and animals. Photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light and dark periods.
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Plants
Many flowering plants (angiosperms) use a photoreceptor protein, such as phytochrome or cryptochrome, 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.
Phytochromes are signaling proteins that promote photomorphogenesis in response to red light and far-red light. There are two forms of phytochromes: red light absorbing, Pr, and far-red light absorbing, Pfr. Pfr, which is the active form of phytochromes, can be reverted to Pr, which is the inactive form, slowly by inducing darkness or by inducing far-red light which makes the process rapid. There are five genes that encode phytochromes in the Arabidopsis thaliana genetic model, PHYA-PHYE. PHYA is the gene that is involved in the regulation of photomorphogenesis in response to far-red light. PHYB is the gene that is involved in regulating photoreversible seed germination in response to red light. PHYC mediates the response between PHYA and PHYB. PHYD and PHYE mediate elongation of the internode and control the time in which the plant flowers.
Cryptochromes are blue-light photoreceptors that regulate responses to blue light. There are two different forms of crytochromes, CRY1 and CRY2, which regulate the inhibition of hypocotyl elongation in response to blue light.
The combination of phytochromes and cryptochromes mediate growth and the flowering of plants in response to red light, far-red light, and blue light.
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 in the northern hemisphere 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). 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:
Some long-day facultative plants are:
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:
Day-neutral plants
Day-neutral plants, such as cucumbers, roses, and tomatoes, 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).
Animal
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 sexual organs.
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.
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). Some mammals are highly seasonal, while humans' seasonality is largely believed to be evolutionary baggage.