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Frost resistance and ice nucleation in leaves of five woody timberline species measured in situ during shoot expansion.

Frost resistance and ice nucleation temperatures of leaves, from bud swelling until after full expansion, were measured in situ for five major woody timberline species with recently developed field freezing equipment. Frost resistance determined in situ on leaves of attached twigs was significantly higher than values determined on detached leaves in laboratory tests (e.g., the temperature at which incipient frost damage was observed (LTi) was 1.2 degrees C higher for detached leaves than for attached leaves of Picea abies (L.) Karst.). Frost resistance of leaves of all species changed significantly during shoot expansion (e.g., changes of 7.2 and 11 degrees C for Rhododendron ferrugineum L. and Larix decidua Mill., respectively). Expanding leaves (between 0 and 60% of full expansion) were the most sensitive to frost, with LTi values ranging from -3.4 degrees C in R. ferrugineum to -6.3 degrees C in L. decidua. Among the studied species, P. abies and R. ferrugineum were the most frost sensitive throughout the shoot elongation period. In situ freezing patterns of leaves of attached twigs also differed from those of leaves of excised twigs. During leaf expansion, two distinct freezing exotherms were always registered in situ. The first freezing event (E1, high-temperature exotherm) was recorded at -1.5 +/- 0.2 degrees C and reflected extracellular ice formation. Exposure of leaves to temperatures at which E1 occurred was, in all cases, noninjurious. The low-temperature exotherm (E2) mostly coincided with frost damage, except for some stages of leaf expansion in R. ferrugineum and P. abies, indicating that in situ freezing exotherms were not accurate estimators of frost damage in these species.

Cold Climate↗

Abiotic factors limiting photosynthesis in Abies lasiocarpa and Picea engelmannii seedlings below and above the alpine timberline.

Most research on the occurrence and stability of alpine timberlines has focused on correlations between adult tree growth and mean temperatures rather than on specific mechanisms. Timberline migration to higher altitude is dependent on new seedling establishment in the tree-line ecotone; however, reductions in photosynthetic carbon gain in establishing seedlings have previously been interpreted solely in terms of decreased seedling survival. Our objective was to evaluate the impact of abiotic factors (temperature, light and water) on photosynthetic carbon gain in young seedlings of the two dominant conifer tree species occurring naturally above (tree-line ecotone site, TS) and below (forest site, FS) a Rocky Mountain timberline in southeastern Wyoming, USA. Coincidentally, measurements were made during an unusually dry summer. Mean daily photosynthesis in seedlings of both Abies lasiocarpa (Hook.) Nutt. (subalpine fir) and Picea engelmannii Parry ex Engelm. (Engelmann spruce) was less at TS than at FS (19 and 29%, respectively). Minimum nighttime temperatures below 2 degrees C were more frequent at TS than at FS and were associated with reduced maximum photosynthesis the following day. Low midday water potentials were associated with a reduction in carbon gain at both sites early in the season, prior to snowmelt, as well as late in the season when soils began to dry. However, the lower photosynthetic rates at TS than at FS appeared to be unrelated to seedling water status because seedlings at both sites had similar xylem pressure potentials. Solar irradiance was highly variable at both sites as a result of uneven shading by neighboring trees, although this variation was substantially reduced on cloudy days (44% of all days observed). Compared with sunny days, cloudy days resulted in greater integrated daily carbon gain at both sites (41% increase at TS and 69% increase at FS), based on a simulated photosynthesis model. Photosynthetic responses to temperature, sunlight and water suggest that variable solar irradiance and nighttime temperatures were major abiotic factors limiting photosynthetic carbon acquisition in these young seedlings, especially for seedlings growing in the tree-line ecotone.

Abies↗

Seasonal variation of soil respiration in a Pinus cembra forest at the upper timberline in the Central Austrian Alps.

Soil respiration (R) of a 95-year-old Pinus cembra L. forest at the alpine timberline was measured continuously from October 2001 to January 2003 with an automated multiplexing gas exchange system. There was significant spatial variability in soil respiration, and R at a soil temperature of 10 degrees C (R10) decreased by about 20% m(-1) with increasing distance from the trunk. Needle litter and fine root density also decreased. The spatially averaged annual soil CO2 efflux was 35 g C m(-2) year(-1) in 2002. About 70% of the temporal variation in soil respiration could be explained by variations in soil temperature, whereas the influence of soil water potential and thus soil water content was negligible because soil water availability was supra-optimal.

Austria↗

Physiological responses of three deciduous conifers (Metasequoia glyptostroboides, Taxodium distichum and Larix laricina) to continuous light: adaptive implications for the early Tertiary polar summer.

Polar regions were covered with extensive forests during the Cretaceous and early Tertiary, and supported trees comparable in size and productivity to those of present-day temperate forests. With a winter of total or near darkness and a summer of continuous, low-angle illumination, these temperate, high-latitude forests were characterized by a light regime without a contemporary counterpart. Although maximum irradiances were much lower than at mid-latitudes, the 24-h photoperiod provided similar integrated light flux. Taxodium, Larix and Metasequoia, three genera of deciduous conifers that occurred in paleoarctic wet forests, have extant, closely related descendents. However, the contemporary relative abundance of these genera differs greatly from that in the paleoarctic. To provide insight into attributes that favor competitive success in a continuous-light environment, we subjected saplings of these genera to a natural photoperiod or a 24-h photoperiod and measured gas exchange, chlorophyll fluorescence, non-structural carbohydrate concentrations, biomass production and carbon allocation. Exposure to continuous light significantly decreased photosynthetic capacity and quantum efficiency of photosystem II in Taxodium and Larix, but had minimal influence in Metasequoia. In midsummer, foliar starch concentration substantially increased in both Taxodium and Larix saplings grown in continuous light, which may have contributed to end-product down-regulation of photosynthetic capacity. In contrast, Metasequoia allocated photosynthate to continuous production of new foliar biomass. This difference in carbon allocation may have provided Metasequoia with a two fold advantage in the paleoarctic by minimizing depression of photosynthetic capacity and increasing photosynthetic surface.

Adaptation, Physiological↗

Models of the spring phenology of boreal and temperate trees: Is there something missing?

According to prevailing theory, air temperature is the main environmental factor regulating the timing of bud burst of boreal and temperate trees. Air temperature has a dual role in this regulation. First, after the cessation of growth in autumn, prolonged exposure to chilling causes rest completion, i.e., removes the physiological growth-arresting conditions inside the bud. After rest completion, prolonged exposure to warm conditions causes ontogenetic development leading to bud burst or flowering. During the past three decades, several simulation models based on chilling and forcing have been developed and tested. In recent modeling studies of the timing of bud burst in mature trees, the simpler thermal-time models that assume forcing starts on a fixed date in the spring have outperformed the chilling-forcing models. We hypothesize that this discrepancy may be due to some element missing from the chilling-forcing models. We tested two new model formulations by introducing reversing, temperature-driven elements that precede forcing and by fitting the models to seven historical time series of data of flowering and leaf bud burst of common boreal tree species. In these tests, both of the new models were generally more accurate in predicting the timing of bud burst than a classical chilling-forcing model, but less accurate than the simple thermal-time model. We therefore conclude that besides chilling, other environmental factors are involved in the regulation of the timing of bud burst. Further work is needed to determine if the regulatory factors derive from air temperature or from some other environmental condition such as changes in light conditions, like day length or night length.

Alnus↗

Responding to winter emergencies.

Nurses need to know how to provide emergency care to victims of wintertime injuries and illnesses--and how to survive such circumstances themselves. In this article, the author, a ski patrol member, describes cold-weather strategies and how to respond to hypothermia, frostbite, winter sports injuries, and avalanches.

Cold Climate↗

Stimulation of tyrosine phosphorylation of a brain protein by hibernation.

Mammalian hibernation is a state of natural tolerance to severely decreased brain blood flow. As protein tyrosine phosphorylation is believed to be involved in the development of resistance to potentially cell-damaging insults, we used immunoblotting for the phosphotyrosine moiety to analyze extracts from various tissues of hibernating and nonhibernating ground squirrels. A single, hibernation-specific phosphoprotein was detected in the brain, but not in any other tissue tested. This protein, designated pp98 to reflect its apparent molecular weight, is distributed throughout the brain, and is associated with the cellular membrane fraction. The presence of the protein is tightly linked to the hibernation state; it is not present in contemporaneously assayed animals that are exposed to the same cold temperature as the hibernators, is present for the duration of a hibernation bout (tested from 1 to 14 days), and disappears within 1 hour of arousal from hibernation. The close association of pp98 with the hibernation state, its presence in cellular membranes, and the known properties of membrane phosphotyrosine proteins suggest that it may transduce a signal for adaptation to the limited availability of oxygen and glucose and low cellular temperature that characterizes hibernation in the ground squirrel.

Animals↗

Trauma in the arctic: an incident report.

On January 29, 1989, a military aircraft crashed on approach to an airfield in Alaska. The subsequent extraction, evacuation, and casualty resuscitation occurred at temperatures lower than -50 degrees F. Both the medical and military literature are replete with instructions and advice regarding the handling of trauma victims in the field in the Arctic environment; however, case reviews of such incidents are rare. In this case, the circumstances of the incident, and the organizations on which it fell, argued for almost ideal handling of the care of the victims. Problems that have an impact on the availability of casualty management, generally, in the Arctic such as a widely scattered population and sparse medical facilities, poorly defined evacuation routes, and marginal communication patterns, were not encountered in this review because of the unusual circumstances of this incident. The difficulties of field care in this environment, even under "ideal" conditions, however, are real and evident in this case history. Transportation assets are critical. Rehearsal, critique, refinement, and re-rehearsal of a mass casualty plan can significantly increase victim survival. Proper protocols and confidence in them take on an importance far beyond their benefit in a more temperate environment. Procedures must be worked out in advance, known by all concerned, and practiced, because the "rescuer" is at the same risk of hypothermia as the "victim" and, in addition, is responsible for thinking for the incapacitated patient. Rote-learned drills will be acknowledged far longer in the cold than unspecific guidelines. Survival for trauma victims may depend on those rote actions.

Accidents, Aviation↗

Effects of reduced ambient temperature on fat utilization during submaximal exercise.

PURPOSE: The influence of cold air exposure on fuel utilization during prolonged cycle exercise was investigated. METHODS: Nine male subjects cycled for 90 min in ambient temperatures of -10 degrees C, 0 degrees C, 10 degrees C, and 20 degrees C. External work performed between conditions was constant. Mean oxygen consumption (VO2) over the 90 min in the 20 degrees C trial corresponded to 64 +/- 5.8% VO2peak. RESULTS: Although mean skin temperature was different between trials (P < 0.05), rectal temperatures were not different. At -10 degrees C and 0 degrees C, the respiratory exchange ratio was higher compared with 10 degrees C and 20 degrees C (0.98 +/- 0.01 and 0.97 +/- 0.01 vs 0.92 +/- 0.01 and 0.91 +/- 0.01; P < 0.05). The associated rates of fat oxidation were lower at -10 degrees C and 0 degrees C compared with 10 degrees C and 20 degrees C (0.15 +/- 0.06 and 0.17 +/- 0.06 vs 0.35 +/- 0.06 and 0.40 +/- 0.04 g.min-1; P < 0.05). Blood glycerol was lower at -10 degrees C and 0 degrees C compared with 20 degrees C (P < 0.05); mean values were 0.13 +/- 0.0, 0.13 +/- 0.0, and 0.18 +/- 0.0 mmol.L-1 for the -10 degrees C, 0 degrees C, and 20 degrees C trials, respectively. Mean VO2 was lower in the -10 degrees C trial than the 20 degrees C trial (2.53 +/- 0.06 vs 2.77 +/- 0.09. L.min-1; P < 0.05). Mean blood glucose concentrations were lower at -10 degrees C than 20 degrees C (4.9 +/- 0.2 vs 5.3 +/- 0.1 mmol.L-1; P < 0.05). Although plasma epinephrine concentrations were greater during the 20 degrees C trial compared with all other trials (P < 0.05), plasma norepinephrine did not differ between trials. CONCLUSION: The diminished fat oxidation at colder temperatures potentially reflects a reduction in lipolysis and/or mobilization of FFA or impairment in the oxidative capacity of the muscle.

Adipose Tissue↗

Arctic and Antarctic exploration including the contributions of physicians and effects of disease in the polar regions.

A history of Arctic and Antarctic exploration, whether to find a Northwest Passage, North Pole, or South Pole, is a story of triumph and tribulation. The hardship experienced by polar explorers in the last 1000 years permeates the tales of achievement. Physicians and surgeons have played prominent roles in all major polar explorations. No significant Arctic voyage, particularly in the last 300 years, was made without a member of the party trained in the management of medical emergencies and in basic surgery. During times of health, surgeons functioned as the voyage naturalists with expertise in biology, botany, zoology, and the writing of scientific catalogs. Spurred by our interest and fascination with the history of polar exploration, we reviewed the roles of physicians and natural scientists in Arctic and Antarctic adventures.

Antarctic Regions↗

Contact lenses in extreme cold environments: response of rabbit corneas.

Contact lenses are worn by many individuals in military and civilian populations. Anecdotal reports have described contact lenses "sticking" and "freezing" to the eye during extreme cold conditions. However, some articles indicate the advantages of wearing contact lenses in cold environments. Military operations frequently taken place in cold regions; therefore, we need to known whether contact lenses can be worn safely in extreme cold. Rabbits were fitted with hard (polymethyl methacrylate) contact lenses and exposed to -28.9 degrees C temperatures with winds up to 78 mph (125 km/hr) for 3-hr periods. The wind-chill factor in these conditions exceeded -67.8 degrees C. No effects of the cold or contact lenses were seen in 85% of the eyes. A few of the eyes, both with contact lenses and without, showed mild superficial fluorescein staining of the cornea which cleared within a few ours after exposure. Histologic examination of the corneas revealed no abnormalities attributable to the cold. Inasmuch as this study showed that rabbits wearing contact lenses in extreme cold suffered no acute deleterious effects to the eyes, the research can be expanded to include human subjects.

Animals↗

Mediterranean Europe as an area of endemism for small mammals rather than a source for northwards postglacial colonization.

There is a general perception that central and northern Europe were colonized by range expansion from Mediterranean refugia at the end of the last glaciation. Data from various species support this scenario, but we question its universality. Our mitochondrial DNA studies on three widespread species of small mammal suggest that colonization may have occurred from glacial refugia in central Europe-western Asia. The haplotypes on the Mediterranean peninsulae are distinctive from those found elsewhere. Rather than contributing to the postglacial colonization of Europe, Mediterranean populations of widespread small mammals may represent long-term isolates undergoing allopatric speciation. This could explain the high endemism of small mammals associated with the Mediterranean peninsulae.

Animals↗