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Cold hardening is the physiological and biochemical process by which an organism prepares for cold weather.
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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. [1] Freezing temperatures induce dehydrative stress on plants, as water absorption in the root and water transport in the plant decreases. [2] 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. [1] Non-acclimatized individuals can survive −5 °C, while an acclimatized individual in the same species can survive −30 °C. Plants that originated in the tropics, like tomato or maize, don't go through cold hardening and are unable to survive freezing temperatures. [3] 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. [2]
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. [2] 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 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. [2]
Cold increases cell membrane permeability [4] and makes the cell shrink, as water is drawn out when ice is formed in the extracellular matrix between cells. [2] To retain the surface area of the cell membrane so it will be able to regain its former volume when temperature rises again, the plant forms more and stronger Hechtian strands. These are tubelike structures that connect the protoplast with the cell wall. When the intracellular water freezes, the cell will expand, and without cold hardening the cell would rupture. To protect the cell membrane from expansion induced damage, the plant cell changes the proportions of almost all lipids in the cell membrane, and increases the amount of total soluble protein and other cryoprotecting molecules, like sugar and proline. [3]
Chilling injury occurs at 0–10 degrees Celsius, as a result of membrane damage, metabolic changes, and toxic buildup. Symptoms include wilting, water soaking, necrosis, chlorosis, ion leakage, and decreased growth. Freezing injury may occur at temperatures below 0 degrees Celsius. Symptoms of extracellular freezing include structural damage, dehydration, and necrosis. If intracellular freezing occurs, it will lead to death. Freezing injury is a result of lost permeability, plasmolysis, and post-thaw cell bursting.
When spring comes, or during a mild spell in winter, plants de-harden, and if the temperature is warm for long enough – their growth resumes. [1]
Cold hardening has also been observed in insects such as the fruit fly and diamondback moth. The insects use rapid cold hardening to protect against cold shock during overwintering periods. [5] [6] Overwintering insects stay awake and active through the winter while non-overwintering insects migrate or die. Rapid cold hardening can be experienced during short periods of undesirable temperatures, such as cold shock in environment temperature, as well as the common cold months. The buildup of cryoprotective compounds is the reason that insects can experience cold hardening. [5] Glycerol is a cryoprotective substance found within these insects capable of overwintering. Through testing, glycerol requires interactions with other cell components within the insect in order to decrease the body's permeability to the cold. [5] When an insect is exposed to these cold temperatures, glycerol rapidly accumulates. Glycerol is known as a non-ionic kosmotrope forming powerful hydrogen bonds with water molecules. The hydrogen bonds in the glycerol compound compete with the weaker bonds between the water molecules causing an interruption in the makeup of ice formation. [7] This chemistry found within the glycerol compound and reaction between water has been used as an antifreeze in the past, and can be seen here when concerning cold hardening. Proteins also play a large role in the cryoprotective compounds that increase ability to survive the cold hardening process and environmental change. Glycogen phosphorylase (GlyP) has been a key protein found during testing to increase in comparison to a controlled group not experiencing the cold hardening. [8] Once warmer temperatures are observed the process of acclimation begins, and the increased glycerol along with other cryoprotective compounds and proteins are also reversed. There is a rapid cold hardening capacity found within certain insects that suggests not all insects can survive a long period of overwintering. Non-diapausing insects can sustain brief temperature shocks but often have a limit to what they can handle before the body can no longer produce enough cryoprotective components.
Inclusive to the cold hardening process being beneficial for insects survival during cold temperatures, it also helps improve the organism's performance. [9] Rapid cold hardening (RCH) is one of the fastest cold temperature responses recorded. [9] This process allows an insect to instantly adapt to the severe weather change without compromising function. The Drosophila melanogaster (common fruit fly) is a frequently experimented insect involving cold hardening. A proven example of RCH enhancing organisms performance comes from courting and mating within the fruit fly. It has been tested that the fruit fly mated more frequently once RCH has commenced in relation to a controlled insect group not experiencing RCH. [9] Most insects experiencing extended cold periods are observed to modify the membrane lipids within the body. Desaturation of fatty acids are the most commonly seen modification to the membrane. [9] When the fruit fly was observed under the stressful climate the survival rate increased in comparison to the fly prior to cold hardening.
In addition to testing on the common fruit fly, Plutella xylostella (diamondback moth) also has been widely studied for its significance in cold hardening. While this insect also shows an increase in glycerol and similar cryoprotective compounds, it also shows an increase in polyols. These compounds are specifically linked to cryoprotective compounds designed to withstand cold hardening. The polyol compound is freeze-susceptible and freeze tolerant. [10] Polyols simply act as a barrier within the insect body by preventing intracellular freezing by restricting the extracellular freezing likely to happen in overwintering periods. [10] During the larval stage of the diamondback moth, the significance of glycerol was tested again for validity. The lab injected the larvae with added glycerol and in turn proved that glycerol is a major factor in survival rate when cold hardening. The cold tolerance is directly proportional to the buildup of glycerol during cold hardening. [10]
Cold hardening of insects improves the survival rate of the species and improves function. Once environmental temperature begins to warm up above freezing, the cold hardening process is reversed and the glycerol and cryoprotective compounds decrease within the body. This also reverts the function of the insect to pre-cold hardening activity.
Abiotic stress is the negative impact of non-living factors on the living organisms in a specific environment. The non-living variable must influence the environment beyond its normal range of variation to adversely affect the population performance or individual physiology of the organism in a significant way.
Freezing is a phase transition where a liquid turns into a solid when its temperature is lowered below its freezing point. In accordance with the internationally established definition, freezing means the solidification phase change of a liquid or the liquid content of a substance, usually due to cooling.
Cryobiology is the branch of biology that studies the effects of low temperatures on living things within Earth's cryosphere or in science. The word cryobiology is derived from the Greek words κρῧος [kryos], "cold", βίος [bios], "life", and λόγος [logos], "word". In practice, cryobiology is the study of biological material or systems at temperatures below normal. Materials or systems studied may include proteins, cells, tissues, organs, or whole organisms. Temperatures may range from moderately hypothermic conditions to cryogenic temperatures.
Supercooling, also known as undercooling, is the process of lowering the temperature of a liquid below its freezing point without it becoming a solid. It is achieved in the absence of a seed crystal or nucleus around which a crystal structure can form. The supercooling of water can be achieved without any special techniques other than chemical demineralization, down to −48.3 °C (−54.9 °F). Droplets of supercooled water often exist in stratus and cumulus clouds. An aircraft flying through such a cloud sees an abrupt crystallization of these droplets, which can result in the formation of ice on the aircraft's wings or blockage of its instruments and probes.
Acclimatization or acclimatisation is the process in which an individual organism adjusts to a change in its environment, allowing it to maintain fitness across a range of environmental conditions. Acclimatization occurs in a short period of time, and within the organism's lifetime. This may be a discrete occurrence or may instead represent part of a periodic cycle, such as a mammal shedding heavy winter fur in favor of a lighter summer coat. Organisms can adjust their morphological, behavioral, physical, and/or biochemical traits in response to changes in their environment. While the capacity to acclimate to novel environments has been well documented in thousands of species, researchers still know very little about how and why organisms acclimate the way that they do.
Antifreeze proteins (AFPs) or ice structuring proteins refer to a class of polypeptides produced by certain animals, plants, fungi and bacteria that permit their survival in temperatures below the freezing point of water. AFPs bind to small ice crystals to inhibit the growth and recrystallization of ice that would otherwise be fatal. There is also increasing evidence that AFPs interact with mammalian cell membranes to protect them from cold damage. This work suggests the involvement of AFPs in cold acclimatization.
An antifreeze is an additive which lowers the freezing point of a water-based liquid. An antifreeze mixture is used to achieve freezing-point depression for cold environments. Common antifreezes also increase the boiling point of the liquid, allowing higher coolant temperature. However, all common antifreeze additives also have lower heat capacities than water, and do reduce water's ability to act as a coolant when added to it.
In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions. It is a physiological state with very specific initiating and inhibiting conditions. The mechanism is a means of surviving predictable, unfavorable environmental conditions, such as temperature extremes, drought, or reduced food availability. Diapause is observed in all the life stages of arthropods, especially insects.
Cytorrhysis is the permanent and irreparable damage to the cell wall after the complete collapse of a plant cell due to the loss of internal positive pressure. Positive pressure within a plant cell is required to maintain the upright structure of the cell wall. Desiccation resulting in cellular collapse occurs when the ability of the plant cell to regulate turgor pressure is compromised by environmental stress. Water continues to diffuse out of the cell after the point of zero turgor pressure, where internal cellular pressure is equal to the external atmospheric pressure, has been reached, generating negative pressure within the cell. That negative pressure pulls the center of the cell inward until the cell wall can no longer withstand the strain. The inward pressure causes the majority of the collapse to occur in the central region of the cell, pushing the organelles within the remaining cytoplasm against the cell walls. Unlike in plasmolysis, the plasma membrane maintains its connections with the cell wall both during and after cellular collapse.
Thermotropism or thermotropic movement is the movement of an organism or a part of an organism in response to heat or changes from the environment's temperature. A common example is the curling of Rhododendron leaves in response to cold temperatures. Mimosa pudica also show thermotropism by the collapsing of leaf petioles leading to the folding of leaflets, when temperature drops.
A cryoprotectant is a substance used to protect biological tissue from freezing damage. Arctic and Antarctic insects, fish and amphibians create cryoprotectants in their bodies to minimize freezing damage during cold winter periods. Cryoprotectants are also used to preserve living materials in the study of biology and to preserve food products.
Belgica antarctica, the Antarctic midge, is a species of flightless midge, endemic to the continent of Antarctica. At 2–6 mm (0.08–0.2 in) long, it is the largest purely terrestrial animal native to the continent. It also has the smallest known insect genome as of 2014, with only 99 million base pairs of nucleotides and about 13500 genes. It is the only insect that can survive year-round in Antarctica.
Insect winter ecology describes the overwinter survival strategies of insects, which are in many respects more similar to those of plants than to many other animals, such as mammals and birds. Unlike those animals, which can generate their own heat internally (endothermic), insects must rely on external sources to provide their heat (ectothermic). Thus, insects persisting in winter weather must tolerate freezing or rely on other mechanisms to avoid freezing. Loss of enzymatic function and eventual freezing due to low temperatures daily threatens the livelihood of these organisms during winter. Not surprisingly, insects have evolved a number of strategies to deal with the rigors of winter temperatures in places where they would otherwise not survive.
Major intrinsic proteins comprise a large superfamily of transmembrane protein channels that are grouped together on the basis of homology. The MIP superfamily includes three subfamilies: aquaporins, aquaglyceroporins and S-aquaporins.
Cryopreservation or cryoconservation is a process where biological material - cells, tissues, or organs - are frozen to preserve the material for an extended period of time. At low temperatures any cell metabolism which might cause damage to the biological material in question is effectively stopped. Cryopreservation is an effective way to transport biological samples over long distances, store samples for prolonged periods of time, and create a bank of samples for users. Molecules, referred to as cryoprotective agents (CPAs), are added to reduce the osmotic shock and physical stresses cells undergo in the freezing process. Some cryoprotective agents used in research are inspired by plants and animals in nature that have unique cold tolerance to survive harsh winters, including: trees, wood frogs, and tardigrades.
Osmoprotectants or compatible solutes are small organic molecules with neutral charge and low toxicity at high concentrations that act as osmolytes and help organisms survive extreme osmotic stress. Osmoprotectants can be placed in three chemical classes: betaines and associated molecules, sugars and polyols, and amino acids. These molecules accumulate in cells and balance the osmotic difference between the cell's surroundings and the cytosol. In plants, their accumulation can increase survival during stresses such as drought. In extreme cases, such as in bdelloid rotifers, tardigrades, brine shrimp, and nematodes, these molecules can allow cells to survive being completely dried out and let them enter a state of suspended animation called cryptobiosis.
Freezing tolerance describes the ability of plants to withstand subzero temperatures through the formation of ice crystals in the xylem and intercellular space, or apoplast, of their cells. Freezing tolerance is enhanced as a gradual adaptation to low temperature through a process known as cold acclimation, which initiates the transition to prepare for subzero temperatures through alterations in rate of metabolism, hormone levels and sugars. Freezing tolerance is rapidly enhanced during the first days of the cold acclimation process when temperature drops. Depending on the plant species, maximum freezing tolerance can be reached after only two weeks of exposure to low temperatures. The ability to control intercellular ice formation during freezing is critical to the survival of freeze-tolerant plants. If intracellular ice forms, it could be lethal to the plant when adhesion between cellular membranes and walls occur. The process of freezing tolerance through cold acclimation is a two-stage mechanism:
The goldenrod gall fly, also known as the goldenrod ball gallmaker, is a species of fly native to North America. The species is best known for the characteristic galls it forms on several species in the Solidago, or goldenrod, genus. The fly's eggs are inserted near the developing buds of the plant. After hatching, the larvae migrate to an area below the plant's developing buds, where they then induce the plant's tissues to form into the hardened, bulbous chamber referred to as a gall. E. solidaginis’s interactions with its host plant(s) and insect, as well as avian, predators have made it the centerpiece of much ecological and evolutionary biology research, and its tolerance of freezing temperatures has inspired studies into the anti-freeze properties of its biochemistry.
Cucujus clavipes is known as the flat bark beetle. It is found throughout North America. These are generally found near tree line under bark of dead poplar and ash trees. C. clavipes are described as phloem-feeding and often predators of other small insects, such as wood-boring beetles, and mites. These are usually seen during spring-summer seasons. Having a cold habitat, these beetles must go through several physiological mechanisms to survive; they are recognised for their ability to change their overwintering mechanisms.
Calcium signaling in Arabidopsis is a calcium mediated signalling pathway that Arabidopsis plants use in order to respond to a stimuli. In this pathway, Ca2+ works as a long range communication ion, allowing for rapid communication throughout the plant. Systemic changes in metabolites such as glucose and sucrose takes a few minutes after the stimulus, but gene transcription occurs within seconds. Because hormones, peptides and RNA travel through the vascular system at lower speeds than the plants response to wounds, indicates that Ca2+ must be involved in the rapid signal propagation. Instead of local communication to nearby cells and tissues, Ca2+ uses mass flow within the vascular system to help with rapid transport throughout the plant. Ca2+ moving through the xylem and phloem acts through a “calcium signature” receptor system in cells where they integrate the signal and respond with the activation of defense genes. These calcium signatures encode information about the stimulus allowing the response of the plant to cater towards the type of stimulus.