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Does leaf position within a canopy affect acclimation of photosynthesis to elevated CO2?. Analysis Of a wheat crop under free-air co2 enrichment

Previous studies of photosynthetic acclimation to elevated CO2 have focused on the most recently expanded, sunlit leaves in the canopy. We examined acclimation in a vertical profile of leaves through a canopy of wheat (Triticum aestivum L.). The crop was grown at an elevated CO2 partial pressure of 55 Pa within a replicated field experiment using free-air CO2 enrichment. Gas exchange was used to estimate in vivo carboxylation capacity and the maximum rate of ribulose-1,5-bisphosphate-limited photosynthesis. Net photosynthetic CO2 uptake was measured for leaves in situ within the canopy. Leaf contents of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), light-harvesting-complex (LHC) proteins, and total N were determined. Elevated CO2 did not affect carboxylation capacity in the most recently expanded leaves but led to a decrease in lower, shaded leaves during grain development. Despite this acclimation, in situ photosynthetic CO2 uptake remained higher under elevated CO2. Acclimation at elevated CO2 was accompanied by decreases in both Rubisco and total leaf N contents and an increase in LHC content. Elevated CO2 led to a larger increase in LHC/Rubisco in lower canopy leaves than in the uppermost leaf. Acclimation of leaf photosynthesis to elevated CO2 therefore depended on both vertical position within the canopy and the developmental stage.

Journal Article↗

Biochemical Changes in Tuber-bearing Solanum Species in Relation to Frost Hardiness during Cold Acclimation.

Biochemical changes in potato leaves during cold acclimation have been examined and compared between a frost-tolerant S. acaule and a frost-susceptible S. tuberosum species. Changes were also examined in S. tuberosum, S. acaule, and S. commersonii species when they were hardened at different temperatures to varying hardiness levels.During three weeks of stepwise cold acclimation, S. acaule increased frost hardiness from -6.0 C (killing temperature) to -9.0 C, whereas frost hardiness of S. tuberosum remained unchanged at -3.0 C. Decreases in DNA content on a dry weight basis in both species suggest that matured leaf cells accumulated more dry matter during acclimation. The advantage of using DNA as a reference for comparing metabolite changes during cold acclimation is discussed.Under the stepwise acclimating conditions, both species showed the same trends for increasing total sugar and starch with an insignificant decrease in leaf water content. High levels of total RNA, rRNA, and total and soluble protein were observed in treated S. acaule plants as compared with controls, but not in S. tuberosum. Levels of total lipid and phospholipid also were high in treated S. acaule plants as compared with controls but decreased in S. tuberosum during acclimation.When S. tuberosum, S. acaule, and S. commersonii potatoes were cold-treated at constant day/night temperatures of 10, 5, and 2 C with 14-hour daylength, each species responds differently in terms of frost hardiness increase upon subjecting plants to a low temperature. For instance, after 20 days at 2 C, a net frost hardiness of 3 and 7 C was observed in S. acaule and S. commersonii, respectively, whereas the frost hardiness in S. tuberosum remained unchanged. Also, various levels of frost hardiness can be achieved in a species by subjecting plants to different low temperature treatments. Under a warm regime of 20/15 C day/night temperatures (14-hour light), both S. acaule and S. commersonii can survive at -4.5 C or colder, whereas S. tuberosum can survive only at -2.5 C.Biochemical changes in the leaf tissue of these species were investigated at 5-day intervals during low temperature treatments. Increases in total sugar and starch were found in all three species during hardening, although S. tuberosum failed to harden. Soluble protein contents were increased in both S. acaule and S. commersonii but decreased in S. tuberosum. RNA contents change in a pattern similar to the soluble protein. Net increases of the soluble proteins were positively and significantly correlated with net increases of frost hardiness in S. acaule and S. commersonii.

Journal Article↗

Influence of cold acclimation on membrane injury in frozen plant tissue.

Cold-acclimated twigs of Amelanchier alnifolia Nutt. released less HCN at -4.5 C than nonacclimated twigs following slow freezing to -25 C or rapid freezing to -78 C. Cold-acclimated twigs frozen slowly to -25 C released more HCN than cold-acclimated twigs frozen only to -4.5 C. Cold-acclimated twigs frozen slowly to -25 C and then rapidly to -78 C released less HCN at -4.5 C than cold-acclimated twigs frozen rapidly to -78 C. In general, K(+) efflux and the inability to reduce triphenyl tetrazolium chloride following freezing and thawing paralleled HCN release at -4.5 C. Because low K(+) efflux and high triphenyl tetrazolium chloride reduction are known to depend upon membrane integrity, the increased K(+) efflux and the decreased triphenyl tetrazolium chloride reduction following freezing and thawing provide indirect evidence that HCN release at -4.5 C is a measure of membrane damage in frozen cells.

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Involvement of Plasma Membrane Alterations in Cold Acclimation of Winter Rye Seedlings (Secale cereale L. cv Puma).

Using plasma membranes with high purity isolated from winter rye seedlings (Secale cereale L. cv Puma) by an aqueous two-polymer phase partition technique, lipid and protein changes were determined during the development of cold hardiness.The degree of fatty acid unsaturation and proportions of phospholipid classes changed only slightly during cold acclimation. A small change, however, was observed in sterol composition; beta-sitosterol increased gradually with a corresponding decrease in campesterol plus stigmasterol during cold acclimation. The phospholipid to protein ratio increased proportionally to the increase in cold hardiness. During the period of acclimation, the sterol to phospholipid ratio declined and was inversely related to the development of cold hardiness.A distinct difference between polypeptide components of nonacclimated and acclimated plasma membrane was observed on two-dimensional sodium dodecyl sulfate slab polyacrylamide gels. A change was also detected in glycopeptides. These results suggest that the plasma membrane is in a dynamic state during cold acclimation, changing in response to the development of cold hardiness.

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Induction of Freezing Tolerance in Spinach during Cold Acclimation.

Spinach (Spinacia oleracea L.) seedlings, grown in soil or on an agar medium in vitro, became cold acclimated when exposed to a constant 5 degrees C. Plants subjected to cold acclimation, beginning 1 week postgermination, attained freezing tolerance levels similar to that achieved by seedlings that were cold acclimated beginning 3 weeks after sowing. Seedlings at 1 week of age had only cotyledonary leaves, while 3-week-old seedlings had developed true leaves. Plants grown in vitro were able to increase in freezing tolerance, but were slightly less hardy than soil-grown plants. These results suggest that spinach, a cool-season crop that begins growth in early spring when subzero temperatures are likely, can undergo cold acclimation at the earliest stages of development following germination. Axenic seedlings, grown in vitro, were used to develop a noninjurious radiolabeling technique. Leaf proteins were radiolabeled to specific activities of 10(5) counts per minute per microgram at 25 degrees C or 5 x 10(4) counts per minute per microgram at 5 degrees C over a 24 hour period. The ability to radiolabel leaf proteins of in vitro grown plants to high specific activities at low temperature, without injury or microbial contamination, will facilitate studies of cold acclimation.

Journal Article↗

Effect of cold acclimation on the incidence of two forms of freezing injury in protoplasts isolated from rye leaves.

The freezing tolerance and incidence of two forms of freezing injury (expansion-induced lysis and loss of osmotic responsiveness) were determined for protoplasts isolated from rye leaves (Secale cereale L. cv Puma) at various times during cold acclimation. During the first 4 weeks of the cold acclimation period, the LT(50) (i.e. the minimum temperature at which 50% of the protoplasts survived) decreased from -5 degrees C to -25 degrees C. In protoplasts isolated from nonacclimated leaves (NA protoplasts), expansion-induced lysis (EIL) was the predominant form of injury at the LT(50). However, after only 1 week of cold acclimation, the incidence of EIL was reduced to less than 10% at any subzero temperature; and loss of osmotic responsiveness was the predominant form of injury, regardless of the freezing temperature. Fusion of either NA protoplasts or protoplasts isolated from leaves of seedlings cold acclimated for 1 week (1-week ACC protoplasts) with liposomes of dilinoleoylphosphatidylcholine also decreased the incidence of EIL to less than 10%. Fusion of protoplasts with dilinoleoylphosphatidylcholine diminished the incidence of loss of osmotic responsiveness, but only in NA protoplasts or 1-week ACC protoplasts that were frozen to temperatures over the range of -5 to -10 degrees C. These results suggest that the cold acclimation process, which results in a quantitative increase in freezing resistance, involves several different qualitative changes in the cryobehavior of the plasma membrane.

Journal Article↗

Microtubules in mesophyll cells of nonacclimated and cold-acclimated spinach : visualization and responses to freezing, low temperature, and dehydration.

Responses of cortical microtubules in spinach (Spinacia oleracea L. cv Bloomsdale) mesophyll cells to freezing, thawing, supercooling, and dehydration were assessed. Microtubules were visualized using a modified procedure for indirect immunofluorescence microscopy. Leaf sections of nonacclimated and cold-acclimated spinach were slowly frozen to various temperatures, fixed while frozen, and microtubules immunolabelled. Both nonacclimated and cold-acclimated cells exhibited nearly complete microtubule depolymerization after ice formation. After 1 hour thawing at 23 degrees C, microtubules in both nonacclimated and cold-acclimated cells repolymerized. With time, however, microtubules in nonacclimated cells again depolymerized. Since microtubules in cells of leaf tissue frozen slowly are subjected to dehydration as well as subzero temperatures, these stresses were applied separately and their effects on microtubules noted. Supercooling induced microtubule depolymerization in both nonacclimated and cold-acclimated cells, but to a smaller extent than did freezing. Exposing leaf sections to solutions of sorbitol (a cell wall-penetrating osmoticum) or polyethylene glycol 10,000 (a nonpenetrating osmoticum) at room temperature caused microtubule depolymerization. The effects of low temperature and dehydration are roughly additive in producing the observed microtubule responses during freezing. Only small differences in microtubule stability were resolved between nonacclimated and cold-acclimated cells.

Journal Article↗

Low-temperature signal transduction: induction of cold acclimation-specific genes of alfalfa by calcium at 25 degrees C.

To study the role of calcium in cold acclimation, we examined the relationship between calcium influx and accumulation of transcripts of two cas (cold acclimation-specific) genes of alfalfa, cas15 and cas18. Whereas a decline in temperature from 25 to 15 degrees C had little effect on the influx of extracellular 45Ca2+, an increasing influx was observed when the temperature was lowered further. The influx of 45Ca2+ at 4 degrees C was nearly 15 times greater than at 25 degrees C. The addition of calcium chelators or of calcium channel blockers, which have been shown to prevent cold acclimation, inhibited the influx of extracellular 45Ca2+ as well as the expression of cas genes at 4 degrees C. The addition of a calcium ionophore or a calcium channel agonist to nonacclimated cells caused the influx of extracellular 45Ca2+ and induced the expression of cas genes at 25 degrees C. These results suggest that a cold-induced calcium influx plays an essential role in cold acclimation. To further study the role of calcium, we isolated two sequences corresponding to calcium-dependent protein kinases. The transcript level of one of them was markedly upregulated at 4 degrees C. We propose a sequence of signaling events that is likely to occur early during cold acclimation and leads to the expression of cas genes and the development of freezing tolerance.

Acclimatization↗

Thermal acclimation of neonates to prolonged cool exposure as regards sleep stages.

The thermal responses of neonates during a cool acclimation period were studied with regard to sleep stages. Sleep stages, body temperatures and metabolic rate (VO2) were studied for seven neonates nursed in incubators and exposed to a cool temperature (thermoneutrality minus 2 degrees C) for 75 h. Each recording session lasted 3 h in the morning: firstly under thermoneutral baseline conditions, then during the first and last 3-h periods of the cool acclimation and finally during the last 3 h of a 24-h recovery period. Sleep structure was modified during the initial hours of cool exposure: the percentage of active sleep increased (AS: +13%, P = 0.028) at the expense of quiet sleep (QS: -11%, P = 0.043). This alteration in sleep structure persisted at the end of the acclimation period. Metabolic heat production only increased in the later period of cool acclimation. Throughout the cool exposure, VO2 increased more (P = 0.040) in QS (+33%) than in AS (+20%) so that by the end of the cool period, VO2 levels were similar in both sleep stages. During cool acclimation, the maintenance of homeothermy is related not only to a change in sleep organization but also to modifications in the thermoregulatory processes in both sleep stages. Considering the importance of AS/QS patterns in the neurobehavioral development of neonates, the present results could have clinical implications for the thermal management of neonates.

Adaptation, Physiological↗

A functional comparison of acclimation to shade and submergence in two terrestrial plant species.

Terrestrial plants experience multiple stresses when they are submerged, caused both by oxygen deficiency due to reduced gas diffusion in water, and by shade due to high turbidity of the floodwater. It has been suggested that responses to submergence are de facto responses to low light intensity. We investigated the extent to which submergence and shade induce similar acclimation responses by comparing two terrestrial Rumex species that differ in their responses to flooding. Our study confirms that there are strong similarities between acclimation responses to shade and submergence. Petiole length, specific leaf area (SLA), chlorophyll parameters and underwater light-compensation points changed at least qualitatively in the same direction. Maximum underwater photosynthesis rate, however, did discriminate between the functionality of the responses, as the acclimation to submergence appeared to be more effective than acclimation to shade at saturating light. We conclude that acclimation to submergence involves more than an increase in SLA to achieve the significant reduction of diffusion resistance for gas exchange between leaves and the water column.

Acclimatization↗

3':5'-Cyclic-AMP phosphodiesterase activities in white and brown adipose tissues of cold-acclimated rats.

3':5'-Cyclic-AMP phosphodiesterase (PDE) (EC 3.1.4.17) activity was measured in interscapular brown adipose tissue (BAT) and in white epididymal adipose tissue of rats acclimated to constant or fluctuating cold. Experiments were carried out on isolated adipocytes or tissue homogenates. In brown or white adipose tissue or isolated adipocyte homogenates, two different apparent Km values were found according to the substrate (cAMP) concentration. The low Km was at about 10(-6) M and the high one at about 10(-4) M. The apparent V of the high Km enzyme was about 10-fold higher than the V of the low Km enzyme. Cold acclimation to constant or fluctuating cold did not modify appreciably the Km or V values. For low substrate concentrations (10(-6)-10(-8) M), the specific activity of PDE expressed per milligram of protein was decreased in BAT adipocytes of the two groups of cold-acclimated rats, compared to controls. Inversely, it was increased in total tissue homogenates. These variations were smaller in fluctuating cold than in constant cold-acclimate rats. They could, in part, induce the increases in lipolysis and in blood flow observed in the BAT of cold-acclimated rats.

3',5'-Cyclic-AMP Phosphodiesterases↗

Catecholamine sensitivity in brown fat cells from cold-acclimated hamsters and rats.

Brown fat cells, freshly isolated from cold-acclimated hamsters and rats, did not respond to norepinephrine addition with the characteristic increase in oxygen consumption (heat production) seen in cells from control animals. However, incubation of these cells for 1 h in a Krebs-Ringer bicarbonate buffer, in the presence of 10 mM pyruvate, fully restored norepinephrine responsiveness. Cells treated in this way from cold-acclimated hamsters (a hibernator) increased the rate of oxygen consumption after maximal norepinephrine stimulation as much as cells from control hamsters; also norepinephrine-stimulated fatty acid release was unaltered, indicating that brown fat cells may partly be responsible for the increase in serum fatty acid level seen during arousal from hibernation. Similarly, preincubated cells from cold-acclimated rats (a nonhibernator) increased oxygen consumption and fatty acid release as much as cells from control rats; this suggests that also in cold-acclimated rats brown fat may supply the circulation with fatty acids during cold stress. Cells from cold-acclimated animals were, however, about 10 times less sensitive to norepinephrine than cells from control animals; this desensitization may be the result of a stimulated phosphodiesterase.

Adaptation, Physiological↗

Proliferation and differentiation of brown adipocytes from interstitial cells during cold acclimation.

The mechanisms of brown adipocyte proliferation and differentiation during cold acclimation (and/or adaptation to hyperphagia) have been studied by quantitative photonic radioautography. [3H]thymidine was injected to warm-acclimated (25 degrees C) rats and to animals exposed to 5 degrees C for 2 days. Samples of interscapular brown adipose tissue were collected for quantitative analysis of mitotic frequencies at various periods of time (4 h-15 days) after the injection of [3H]thymidine, the rats being maintained at the temperatures to which they were initially exposed. Confirming our previous results [Bukowiecki et al., Am. J. Physiol. 242 (Endocrinol. Metab. 5): E353-E359, 1982], it was found that cold exposure for 2 days markedly enhanced mitotic activity in endothelial cells, interstitial cells, and brown preadipocytes rather than in fully differentiated brown adipocytes. The total tissue labeling index (percent of labeled nuclei) increased approximately 70 times over control values. We now report that cellular labeling progressively increased in mature brown adipocytes during cold acclimation, whereas it correspondingly decreased in interstitial cells and brown preadipocytes. This indicates that the sequence of events for cellular differentiation is interstitial cells----brown preadipocytes----mature brown adipocytes. Remarkable, labeling frequency did not change in endothelial cells during cold acclimation demonstrating that these cells cannot be considered as progenitors of brown adipocytes. It is suggested that brown adipocyte proliferation and differentiation from interstitial cells represent the fundamental phenomena explaining the enhanced capacity of cold-acclimated and/or hyperphagic rats to respond calorigenically to catecholamines.

Adaptation, Physiological↗

Cold acclimation induces desensitization to adenosine in brown fat cells without changing receptor binding.

The ability of brown fat cells isolated from control and cold-acclimated hamsters to respond to adenosine was investigated. In measurements of the rate of oxygen consumption, it was observed that cells from control hamsters responded as expected to addition of adenosine deaminase, 3-isobutyl-1-methylxanthine (IBMX), or 2-chloroadenosine (i.e., norepinephrine dose-response curves were shifted to left in presence of adenosine deaminase or IBMX and to right with 2-chloroadenosine). However, brown fat cells isolated from cold-acclimated hamsters, under identical conditions, showed almost complete absence of adenosine control. Thus acclimation to cold induced a desensitization to adenosine by physiological means. To evaluate the molecular mechanism underlying desensitization to adenosine, [3H]phenylisopropyladenosine ([3H]PIA) binding to brown fat membranes from control and cold-acclimated hamsters was investigated. [3H]PIA bound with similar high affinity (KD approximately 5 nM) and saturability (Bmax approximately 15 fmol/mg protein) in both membrane preparations, demonstrating that desensitization to adenosine was not due to changes in adenosine receptor number or receptor affinity for adenosine. Furthermore, GTP induced a reduction in [3H]PIA affinity in brown fat membranes from both control and cold-acclimated hamsters, indicating that desensitization was probably not due to an uncoupling between the receptor and Gi protein. It was therefore concluded that the adenosine desensitization process may be located at the Gi protein-adenylate cyclase interaction.

1-Methyl-3-isobutylxanthine↗

Alterations in adipocyte response to lipolytic hormones during cold acclimation.

The effects of cold exposure (7 days, 5 degrees C) and cold acclimation (21 days, 5 degrees C) on the regulation of lipolysis were investigated in adipocytes isolated from epididymal fat pads of rats. Catecholamines stimulated lipolysis in an affinity sequence typical of the beta 1-adrenoceptor subtype: one-half maximum velocity (1/2 Vmax) isoproterenol (35 nM) much greater than 1/2 Vmax norepinephrine (150 nM) approximately 1/2 Vmax epinephrine (200 nM). Cold exposure markedly decreased the sensitivity (1/2 Vmax) and the responsiveness (Vmax) of the adipocytes to the lipolytic action of catecholamines. Addition of adenosine deaminase to fat cells isolated from cold-exposed rats did not normalize the lipolytic activity, suggesting that extracellular adenosine was not responsible for the obtunded lipolysis. This effect of cold exposure was transient as the lipolytic response to catecholamines was normal in fully cold-acclimated animals. Remarkably, the responsiveness of adipocytes to the lipolytic action of glucagon (200 nM) and adrenocorticotropic hormone (ACTH, 1 microM) progressively increased during cold acclimation. Adipocyte lipolytic response to dibutyryl adenosine 3',5'-cyclic monophosphate (cAMP) and theophylline was normal in cold-exposed rats, indicating that the lipolytic defect resides at an early step in the lipolytic cascade (pre-cAMP). On the other hand, the antilipolytic effect of insulin on norepinephrine-induced lipolysis significantly decreased during cold acclimation, particularly at physiological levels of insulin (nanomolar level). These results demonstrate that the transient decrease in the lipolytic action of catecholamines observed during cold acclimation is compensated by 1) an increased responsiveness of adipocytes to glucagon and ACTH and 2) by a decreased effectiveness of insulin to induce antilipolysis.(ABSTRACT TRUNCATED AT 250 WORDS)

Acclimatization↗

Heat acclimation and physical training adaptations of young women using different contraceptive hormones.

Although endogenous and exogenous steroid hormones affect numerous physiological processes, the interactions of reproductive hormones, chronic exercise training, and heat acclimation are unknown. This investigation evaluated the responses and adaptations of 36 inactive females [age 21 +/- 3 (SD) yr] as they undertook a 7- to 8-wk program [heat acclimation and physical training (HAPT)] of indoor heat acclimation (90 min/day, 3 days/wk) and outdoor physical training (3 days/wk) while using either an oral estradiol-progestin contraceptive (ORAL, n = 15), a contraceptive injection of depot medroxyprogesterone acetate (DEPO, n = 7), or no contraceptive (EU-OV, n = 14; control). Standardized physical fitness and exercise-heat tolerance tests (36.5 degrees C, 37% relative humidity), administered before and after HAPT, demonstrated that the three subject groups successfully (P < 0.05) acclimated to heat (i.e., rectal temperature, heart rate) and improved muscular endurance (i.e., sit-ups, push-ups, 4.6-km run time) and body composition characteristics. The stress of HAPT did not disrupt the menstrual cycle length/phase characteristics, ovulation, or plasma hormone concentrations of EU-OV. No between-group differences (P > 0.05) existed for rectal and skin temperatures or metabolic, cardiorespiratory, muscular endurance, or body composition variables. A significant difference post-HAPT in the onset temperature of local sweating, ORAL (37.2 +/- 0.4 degrees C) vs. DEPO (37.7 +/- 0.2 degrees C), suggested that steroid hormones influenced this adaptation. In summary, virtually all adaptations of ORAL and DEPO were similar to EU-OV, suggesting that exogenous reproductive hormones neither enhanced nor impaired the ability of women to complete 7-8 wk of strenuous physical training and heat acclimation.

Acclimatization↗

Thyroid cold acclimation influences on norepinephrine metabolism in brown fat.

Norepinephrine turnover rates and tyrosine hydroxylase activities were determined in the interscapular brown fat pad of the rat during cold acclimation, hyperthyroxinism, and after thyroidectomy. Rats were cold acclimated by placement in a cold room, one rat to a cage, for a period of 6 wk. Hyperthyroxinism was induced by daily subcutaneous injections of L-thyroxine (1 mg/kg) for 6 days. Norepinephrine turnover rate and enzyme activity were determined at the end of each experimental period and at 8 wk after thyroidectomy. The rate of norepinephrine turnover increased during cold acclimation and hyperthyroxinism and decreased after thyroidectomy. Cold acclimation resulted in a significant increase in tyrosine hydroxylase activity, whereas no significant effect on enzyme activity was observed in hyperthyroxinism or after thyroidectomy. None of the conditions produced a change compared to controls in the apparent Km of tyrosine hydroxylase for L-tyrosine. Cold acclimation resulted in a significant decrease in the apparent Km of tyrosine hydroxylase for pterin cofactor, whereas thyroxine treatment and thyroidectomy had no effect.

Acclimatization↗

Thyroid function and cold acclimation in the hamster, Mesocricetus auratus.

Basal metabolic rate (BMR), thyroxine utilization rate (T4U), and triiodothyronine utilization rate (T3U) were measured in cold-acclimated (CA) and room temperature-acclimated (RA) male golden hamsters, Mesocricetus auratus. Hormone utilization rates were calculated via the plasma disappearance technique using 125I-labeled hormones and measuring serum hormone levels via radioimmunoassay. BMR showed a significant 28% increase with cold acclimation from 4.50 +/- 0.05 to 5.77 +/- 0.10 ml O2 X h-1 X g-2/3. The same cold exposure also produced a 32% increase in T4U (10.75 +/- 0.51 vs. 14.19 +/- 0.75 ng X day-1 X g-2/3), and a 204% increase in T3U (5.51 +/- 0.53 vs. 16.77 +/- 1.35). The much greater increase in T3U implies that previous assessments of the relationship between cold acclimation and thyroid function may have been underestimated and that cold exposure induces both quantitative and qualitative changes in thyroid function. It is concluded that in the cold-acclimated state, T3U more accurately reflects thyroid function than does T4U. A mechanism for the cold-induced change in BMR is proposed, for which alterations in four aspects of thyroid function are required: a decrease in plasma T4 binding, an elevation of the pituitary T4 "set point," a preferential shift in deiodinase activity from reverse T3 to T3 production, and an increase in the thyroidal secretion of T3.

Acclimatization↗