Cold hardening

Cold hardening is the physiological and biochemical process by which an organism prepares for cold weather.

Plants in temperate and polar regions adapt to winter and sub zero temperatures by relocating nutrients from leaves and shoots to storage organs. Freezing temperatures induce dehydrative stress on plants, as water absorption in the root and water transport in the plant decreases. Water in and between cells in the plant freezes and expands, causing tissue damage. Cold hardening is a process in which a plant undergoes physiological changes to avoid, or mitigate cellular injuries caused by sub-zero temperatures. Non-acclimatized individuals can survive -5 °C, while a acclimatized individual in the same species can survive -30°. Plants that originated in the tropics, like tomato or maize, don’t go through cold hardening and are unable to survive freezing temperatures. The plant starts the adaptation by exposure to cold yet still not freezing temperatures. The process can be divided into three steps. First the plant perceives low temperature, then converts the signal to activate or repress expression of appropriate genes. Finally, it uses these genes to combat the stress, caused by sub-zero temperatures, affecting its living cells. Many of the genes and responses to low temperature stress are shared with other abiotic stresses, like drought or salinity.

When temperature drops, the membrane fluidity, RNA and DNA stability, and enzyme activity change. These, in turn, affect transcription, translation, intermediate metabolism, and photosynthesis, leading to an energy imbalance. This energy imbalance is thought to be one of the ways the plant detects low temperature. Experiments on arabidopsis show that the plant detects the change in temperature, rather than the absolute temperature. The rate of temperature drop is directly connected to the magnitude of calcium influx, from the space between cells, into the cell. Calcium channels in the cell membrane detect the temperature drop, and promotes expression of low temperature responsible genes in alfalfa and arabidopsis. The response to the change in calcium elevation depends on the cell type and stress history. Shoot tissue will respond more than a root cells, and a cell that already is adapted to cold stress will respond more than one that has not been through cold hardening before. Light doesn’t control the onset of cold hardening directly, but shortening of daylight is associated with fall, and starts production of reactive oxygen species and excitation of photosystem 2, which influences low-temp signal transduction mechanisms. Plants with compromised perception of day length have compromised cold acclimation.

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