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PubMed2000

Stable isotopes examined across a migratory divide in Scandinavian willow warblers (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) reflect their African winter quarters.

The C and N isotopes of feathers from two subspecies of willow warblers Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) are isotopically distinct. Our analysis of 138 adult males from 14 sites distributed across Sweden shows that the mean delta15N and delta13C values of subspecies acredula (from latitudes above 63 degrees N) were significantly higher than the mean delta15N and delta13C values of subspecies trochilus (from latitudes below 61 degrees N). The analysed willow warbler feathers had been moulted in the winter quarters and the observed isotopic signatures should thus reflect the isotopic pattern of food assimilated in Africa. The isotopic data observed in Sweden match the cline in morphology, both showing abrupt changes around 62 degrees N. This result agrees with data from ringing recoveries indicating that the two subspecies occupy geographically and isotopically distinct wintering grounds in Africa. Our isotopic data suggest that analysis of stable isotopes of C and N is a promising method to track wintering quarters of European birds that migrate to Africa.

Africa↗
PubMed2004

PAF1-complex-mediated histone methylation of FLOWERING LOCUS C chromatin is required for the vernalization-responsive, winter-annual habit in Arabidopsis.

The winter-annual habit (which typically involves a requirement for exposure to the cold of winter to flower in the spring) in Arabidopsis thaliana is mainly due to the repression of flowering by relatively high levels of FLC expression. Exposure to prolonged cold attenuates FLC expression through a process known as vernalization and thus permits flowering to occur in the spring. Here we show that the elevated FLC expression characteristic of nonvernalized winter annuals requires two genes, EARLY FLOWERING 7 (ELF7) and EARLY FLOWERING 8 (ELF8), that are homologs of components of the PAF1 complex of Saccharomyces cerevisiae. Furthermore, ELF7 and ELF8 are also required for the expression of other genes in the FLC clade of flowering repressors such as MAF2 and FLM. FLC, FLM, and MAF2 are involved in multiple flowering pathways that account for the broad effects of elf7 and elf8 mutations on flowering behavior. ELF7 and ELF8 are required for the enhancement of histone 3 trimethylation at Lys 4 in FLC chromatin. This modification of FLC chromatin appears to be required to elevate FLC expression to levels that can delay flowering in plants that have not been vernalized. A model of the role of ELF7, ELF8, and other previously described genes in the modification of the chromatin of flowering repressors is presented.

Acclimatization↗
PubMed1992

Antifreeze protein produced endogenously in winter rye leaves.

After cold acclimation, winter rye (Secale cereale L.) is able to withstand the formation of extracellular ice at freezing temperatures. We now show, for the first time, that cold-acclimated winter rye plants contain endogenously produced antifreeze protein. The protein was extracted from the apoplast of winter rye leaves, where ice forms during freezing. After partial purification, the protein was identified as antifreeze protein because it modified the normal growth pattern of ice crystals and depressed the freezing temperature of water noncolligatively.

Journal Article↗
PubMed1994

Characterization and Quantification of Intrinsic Ice Nucleators in Winter Rye (Secale cereale) Leaves.

Extracellular ice formation in frost-tolerant organisms is often initiated at specific sites by ice nucleators. In this study, we examined ice nucleation activity (INA) in the frost-tolerant plant winter rye (Secale cereale). Plants were grown at 20[deg]C, at 5[deg]C with a long day, and at 5[deg]C with a short day (5[deg]C-SD). The threshold temperature for INA was -5 to -12[deg]C in winter rye leaves from all three growth treatments. Epiphytic ice nucleation-active bacteria could not account for INA observed in the leaves. Therefore, the INA must have been produced endogenously. Intrinsic rye ice nucleators were quantified and characterized using single mesophyll cell suspensions obtained by pectolytic degradation of the leaves. The most active ice nucleators in mesophyll cell suspensions exhibited a threshold ice nucleation temperature of -7[deg]C and occurred infrequently at the rate of one nucleator per 105 cells. Rye cells were treated with chemicals and enzymes to characterize the ice nucleators, which proved to be complexes of proteins, carbohydrates, and phospholipids, in which both disulfide bonds and free sulfhydryl groups were important for activity. Carbohydrates and phospholipids were important components of ice nucleators derived from 20[deg]C leaves, whereas the protein component was more important in 5[deg]C-SD leaves. This difference in composition or structure of the ice nucleators, combined with a tendency for more frequent INA, suggests that more ice nucleators are produced in 5[deg]C-SD leaves. These additional ice nucleators may be a component of the mechanism for freezing tolerance observed in winter rye.

Journal Article↗
PubMed1994

Effects of a Short-Term Shift to Low Temperature and of Long-Term Cold Hardening on Photosynthesis and Ribulose-1,5-Bisphosphate Carboxylase/Oxygenase and Sucrose Phosphate Synthase Activity in Leaves of Winter Rye (Secale cereale L.).

The effect of a short-term (hours) shift to low temperature (5[deg]C) and long-term (months) cold hardening on photosynthesis and carbon metabolism was studied in winter rye (Secale cereale L. cv Musketeer). Cold-hardened plants grown at 5[deg]C exhibited 25% higher in situ CO2 exchange rates than nonhardened plants grown at 24[deg]C. Cold-hardened plants maintained these high rates throughout the day, in contrast to nonhardened plants, which showed a gradual decline in photosynthesis after 3 h. Associated with the increase in photosynthetic capacity following cold hardening was an increase in ribulose-1,5-bisphosphate carboxylase/oxygenase and sucrose phosphate synthase activity and 3- to 4-fold increases in the pools of associated metabolites. Leaves of nonhardened plants shifted overnight to 5[deg]C required 9 h in the light at 5[deg]C before maximum rates of photosynthesis were reached. The gradual increase in photosynthesis in leaves shifted to 5[deg]C was correlated with a sharp decline in the 3-phosphoglycerate/triose phosphate ratio and by an increase in the ribulose bisphosphate/3-phosphoglycerate ratio, indicating the gradual easing of aninorganic phosphate-mediated feedback inhibition on photo-synthesis. We suggest that the strong recovery of photosynthesis in winter rye following cold hardening indicates that the buildup of photosynthetic enzymes, as well as those involved in sucrose synthesis, is an adaptive response that enables these plants to maximize the production of sugars that have both cryoprotective and storage functions that are critical to the performance of these cultivars during over-wintering.

Journal Article↗
PubMed1999

Winter survival of transgenic alfalfa overexpressing superoxide dismutase

To test the hypothesis that enhanced tolerance of oxidative stress would improve winter survival, two clones of alfalfa (Medicago sativa) were transformed with a Mn-superoxide dismutase (Mn-SOD) targeted to the mitochondria or to the chloroplast. Although Mn-SOD activity increased in most primary transgenic plants, both cytosolic and chloroplastic forms of Cu/Zn-SOD had lower activity in the chloroplast SOD transgenic plants than in the nontransgenic plants. In a field trial at Elora, Ontario, Canada, the survival and yield of 33 primary transgenic and control plants were compared. After one winter most transgenic plants had higher survival rates than control plants, with some at 100%. Similarly, some independent transgenic plants had twice the herbage yield of the control plants. Prescreening the transgenic plants for SOD activity, vigor, or freezing tolerance in the greenhouse was not effective in identifying individual transgenic plants with improved field performance. Freezing injury to leaf blades and fibrous roots, measured by electrolyte leakage from greenhouse-grown acclimated plants, indicated that the most tolerant were only 1 degrees C more freezing-tolerant than alfalfa clone N4. There were no differences among transgenic and control plants for tetrazolium staining of field-grown plants at any freezing temperature. Therefore, although many of the transgenic plants had higher winter survival rates and herbage yield, there was no apparent difference in primary freezing injury, and therefore, the trait is not associated with a change in the primary site of freezing injury.

Journal Article↗
PubMed2001

Ethylene induces antifreeze activity in winter rye leaves.

Antifreeze activity is induced by cold temperatures in winter rye (Secale cereale) leaves. The activity arises from six antifreeze proteins that accumulate in the apoplast of winter rye leaves during cold acclimation. The individual antifreeze proteins are similar to pathogenesis-related proteins, including glucanases, chitinases, and thaumatin-like proteins. The objective of this study was to study the regulation of antifreeze activity in response to ethylene and salicyclic acid, which are known regulators of pathogenesis-related proteins induced by pathogens. Nonacclimated plants treated with salicylic acid accumulated apoplastic proteins with no antifreeze activity. In contrast, when nonacclimated plants were exposed to ethylene, both antifreeze activity and the concentration of apoplastic protein increased in rye leaves. Immunoblotting revealed that six of the seven accumulated apoplastic proteins consisted of two glucanases, two chitinases, and two thaumatin-like proteins. The ethylene-releasing agent ethephon and the ethylene precursor 1-aminocyclopropane-1-carboxylate also induced high levels of antifreeze activity at 20 degrees C, and this effect could be blocked by the ethylene inhibitor AgNO(3). When intact rye plants were exposed to 5 degrees C, endogenous ethylene production and antifreeze activity were detected within 12 and 48 h of exposure to cold, respectively. Rye plants exposed to drought produced both ethylene and antifreeze activity within 24 h. We conclude that ethylene is involved in regulating antifreeze activity in winter rye in response to cold and drought.

Antifreeze Proteins↗
PubMed1979

Protoplasts surviving freezing to -196 C and osmotic dehydration in 5 molar salt solutions prepared from the bark of winter black locust trees.

Free protoplasts were prepared from the living bark tissue of the trunk of summer and winter black locust trees by enzymic digestion of thin slices of the tissue for 3 hours in a medium containing 2% Onozuka cellulase, 2% Rhozyme pectinase, and 2% Driselase in mannitol solutions using 0.4 molar mannitol for summer tissue and 1.0 molar mannitol for winter tissues. Cleaned suspensions of protoplasts and also thin slices of tissue with cells intact were frozen to temperatures of -10 C, -20 C, -30 C, -40 C and liquid nitrogen in sucrose and balanced salt solutions. Similar suspensions of protoplasts were also subjected to strong osmotic dehydration (plasmorrhysis) in a series of balanced salt solutions of increasing molarity. Tests for survival showed that protoplasts retain the same properties of either extreme susceptibility or extreme resistance to injury by freezing or osmotic dehydration as the cells from which they are prepared. Winter protoplasts showed capability for tolerating freezing to -196 C and plasmorrhysis in 5 molar salt solutions. These results indicate that protoplasts are a valid and useful system for investigating the properties of the protoplasm and surface membranes associated with the seasonal development of extreme hardiness in the cells of woody plants.

Journal Article↗
PubMed1981

Deep undercooling of tissue water and winter hardiness limitations in timberline flora.

Deep undercooled tissue water, which froze near -40 C, was found in winter collected stem and leaf tissue of the dominant timberline tree species of the Colorado Rocky Mountains, Engelmann spruce (Picea engelmannii (Parry) Engelm.) and subalpine fir (Abies lasiocarpa (Hook.) Nutt.), and in numerous other woody species in and below the subalpine vegetation zone. Previous work on numerous woody plants indicates that deep undercooling in xylem makes probable a -40 C winter hardiness limit in stem tissue. Visual injury determinations and electrolyte loss measurements on stem tissue revealed injury near -40 C associated with the freezing of the deep undercooled stem tissue water. These results suggest that the winter hardiness limit of this woody flora is near -40 C. The relevance of deep undercooling in relation to timberline, the upper elevational limit of the subalpine forest, is discussed.

Journal Article↗
PubMed1985

Regulation of RNA Synthesis by DNA-Dependent RNA Polymerases and RNases during Cold Acclimation in Winter and Spring Wheat.

Chromatin DNA-dependent RNA polymerases and RNases activities were measured in winter and spring varieties to understand the overall regulation of RNA synthesis during cold acclimation. We found that total RNA polymerase activities were significantly higher in chromatin isolated from winter wheat compared to the spring wheat during the acclimation period. This increase was parallel to the increase in protein and RNA contents during hardening. The ratio of RNA polymerase I to RNA polymerase II activity was higher than 2 in winter wheat after 30 days of hardening compared, to a ratio of 0.90 under the nonhardening conditions. The increase in activity and the ratio of polymerase I to polymerase II was maintained after the separation of the enzymes from the template, suggesting that RNA synthesis is regulated in part at the enzyme level. On the other hand, the chromatin associated RNase activity decreased in both varieties during acclimation, indicating a nonspecific inhibition caused by low temperature rather than a selective genetic response associated with cold acclimation.

Journal Article↗
PubMed1991

Low growth temperature effects a differential inhibition of photosynthesis in spring and winter wheat.

In vivo room temperature chlorophyll a fluorescence coupled with CO(2) and O(2) exchange was measured to determine photosynthetic limitation(s) for spring and winter wheat (Triticum aestivum L.) grown at cold-hardening temperatures (5 degrees C/5 degrees C, day/night). Plants of comparable physiological stage, but grown at nonhardening temperatures (20 degrees C/16 degrees C, day/night) were used in comparison. Winter wheat cultivars grown at 5 degrees C had light-saturated rates of CO(2) exchange and apparent photon yields for CO(2) exchange and O(2) evolution that were equal to or greater than those of winter cultivars grown at 20 degrees C. In contrast, spring wheat cultivars grown at 5 degrees C showed 35% lower apparent photon yields for CO(2) exchange and 25% lower light-saturated rates of CO(2) exchange compared to 20 degrees C grown controls. The lower CO(2) exchange capacity is not associated with a lower efficiency of photosystem II activity measured as either the apparent photon yield for O(2) evolution, the ratio of variable to maximal fluorescence, or the level of reduced primary quinone electron acceptor maintained at steady-state photosynthesis, and is most likely associated with carbon metabolism. The lower CO(2) exchange capacity of the spring cultivars developed following long-term exposure to low temperature and did not occur following over-night exposure of nonhardened plants to 5 degrees C.

Journal Article↗
PubMed2005

Establishment of the vernalization-responsive, winter-annual habit in Arabidopsis requires a putative histone H3 methyl transferase.

Winter-annual accessions of Arabidopsis thaliana are often characterized by a requirement for exposure to the cold of winter to initiate flowering in the spring. The block to flowering prior to cold exposure is due to high levels of the flowering repressor FLOWERING LOCUS C (FLC). Exposure to cold promotes flowering through a process known as vernalization that epigenetically represses FLC expression. Rapid-cycling accessions typically have low levels of FLC expression and therefore do not require vernalization. A screen for mutants in which a winter-annual Arabidopsis is converted to a rapid-cycling type has identified a putative histone H3 methyl transferase that is required for FLC expression. Lesions in this methyl transferase, EARLY FLOWERING IN SHORT DAYS (EFS), result in reduced levels of histone H3 Lys 4 trimethylation in FLC chromatin. EFS is also required for expression of other genes in the FLC clade, such as MADS AFFECTING FLOWERING2 and FLOWERING LOCUS M. The requirement for EFS to permit expression of several FLC clade genes accounts for the ability of efs lesions to suppress delayed flowering due to the presence of FRIGIDA, autonomous pathway mutations, or growth in noninductive photoperiods. efs mutants exhibit pleiotropic phenotypes, indicating that the role of EFS is not limited to the regulation of flowering time.

Alleles↗
PubMed2005

High prevalence of self-reported winter depression in a Swedish county.

The prevalence of winter depression was unknown in Sweden, therefore prevalence figures of seasonal affective disorder (SAD) and subsyndromal SAD (S-SAD) were estimated. Age and gender differences, prevalence in the group of non-responders and some psychometric qualities of the Seasonal Pattern Assessment Questionnaire (SPAQ) were calculated. A modified version of the SPAQ was sent to a random sample of 2500 persons (response rate 66.3%, n=1657) between 18 and 64 years residing in Dalarna, a county in central Sweden. The sample was proportionally stratified according to age, gender and home municipality. The prevalence of winter SAD was estimated at 8% and S-SAD at 10.8%. It was approximately twice as common among women and younger persons. A total of 3.1% reported seasonal problems to be severe or disabling and 19.3% that everyday life was negatively affected. Experiencing seasonally changing depressive symptoms was common in the population. Factor analysis of the Global Seasonal Score resulted in one factor and the internal consistency was 0.88 (Cronbach's alpha). The results indicate that self-reported recurrent depression during winter is common in Sweden and should therefore receive more attention from health care authorities.

Adolescent↗
PubMed2006

Modelled and measured dynamics of viruses in Arctic winter sea-ice brines.

We describe a model based on diffusion theory and the temperature-dependent mechanism of brine concentration in sea ice to argue that, if viruses partition with bacteria into sea-ice brine inclusions, contact rates between the two can be higher in winter sea ice than in seawater, increasing the probability of infection and possible virus production. To examine this hypothesis, we determined viral and bacterial concentrations in select winter sea-ice horizons using epifluorescence microscopy. Viral concentrations ranged from 1.6 to 82 x 10(6) ml(-1) of brine volume of the ice, with highest values in brines from coldest (-24 to -31 degrees C) ice horizons. Calculated virus-bacteria contact rates in underlying -1 degrees C seawater were similar to those in brines of -11 degrees C ice but up to 600 times lower than those in ice brines at or below -24 degrees C. We then incubated native bacterial and viral assemblages from winter sea ice for 8 days in brine at a temperature (-12 degrees C) and salinity ( approximately 160 psu) near expected in situ values, monitoring their concentrations microscopically. While different cores yielded different results, consistent with known spatial heterogeneity in sea ice, these experiments provided unambiguous evidence for viral persistence and production, as well as for bacterial growth, in -12 degrees C brine.

Arctic Regions↗
PubMed2005

Summer and winter sensitivity of leaves and xylem to minimum freezing temperatures: a comparison of co-occurring Mediterranean oaks that differ in leaf lifespan.

Freezing sensitivity of leaves and xylem was examined in four co-occurring Mediterranean oaks (Quercus spp.) grown in a common garden to determine whether freezing responses of leaves and xylem were coordinated and could be predicted by leaf lifespan. Freezing-induced embolism and loss of photosynthetic function were measured after overnight exposure to a range of subzero temperatures in both summer and winter. Both measures were found to be dependent on minimum freezing temperature and were correlated with leaf lifespan and vessel diameter. The dependence of xylem embolism on minimum freezing temperature may result from the decline in water potential with ice temperature that influences the redistribution of water during freezing and leads to an increase in xylem tension. Winter acclimatization had a relatively small effect on the vulnerability to freezing-induced embolism, although leaf photosynthetic function showed a strong acclimatization response, particularly in the two evergreen species. Quercus ilex, the species with the longest leaf lifespan and narrowest vessel diameters, showed the highest freezing tolerance. This helps explain its ability to inhabit a broad range throughout the Mediterranean region. By contrast, the inability of the deciduous oaks to maintain photosynthetic and vascular function throughout the winter indicates a competitive disadvantage that may prevent them from expanding their ranges.

Climate↗
PubMed2006

Winter down-regulation of intrinsic photosynthetic capacity coupled with up-regulation of Elip-like proteins and persistent energy dissipation in a subalpine forest.

Overwintering, sun-exposed and photosynthetically inactive evergreens require powerful photoprotection. The goal of this study was to seasonally characterize photosynthesis and key proteins/components involved in electron transport and photoprotection. Maximal photosystem II (PSII) efficiency and photosynthetic capacity, amounts of zeaxanthin (Z), antheraxanthin (A), pheophytin and proteins (oxygen-evolving 33 kDa protein (OEC), PSII core protein D1 and subunit S (PsbS) protein, and members of the early light-inducible protein (Elip) family) were assessed in five conifer species at high altitude and in ponderosa pine (Pinus ponderosa) at moderate altitude during summer and winter. Relative to summer, winter down-regulation of photosynthetic capacity and loss of PSII efficiency at the high-altitude sites were paralleled by decreases in OEC, D1, and pheophytin; massive nocturnal retention of (Z + A) and up-regulation of two to four proteins cross-reactive with anti-Elip antibodies; and no change in PsbS amount. By contrast, ponderosa pine at moderate altitude exhibited no down-regulation of photosynthetic capacity, smaller depressions in PSII efficiency, and less up-regulation of Elip family members. These results support a function for members of the Elip family in the acclimation of sun-exposed needles that down-regulate photosynthesis during winter. A possible role in sustained photoprotection is considered.

Altitude↗
PubMed1973

Electrical properties of toad sartorius muscle fibres in summer and winter.

1. The area and circumference of surface fibres of sartorius muscles were measured from photomicrographs of frozen sections of whole muscles, and compared with the values obtained assuming a circular cross-section. The latter assumption gave an over-estimate of the mean area of 28%, but only a 2% over-estimate of the circumference. In isolated, single fibres, the assumption gave over-estimates of 25 and 6%, of area and circumference respectively.2. The passive electrical properties of fibres were different in summer and winter. The mean internal resistivity, membrane resistance and membrane capacitance were 147 Omega.cm, 7.6 kOmega.cm(2) and 4 muF/cm(2) in summer, and 194 Omega.cm, 3.9 kOmega.cm(2) and 6.7 muF/cm(2) in winter, in fibres of comparable diameters in situ. In single fibres in summer, the mean values were 120 Omega.cm, 8.6 kOmega.cm(2) and 3.6 muF/cm(2).3. In glycerol-treated fibres the mean specific membrane capacitance was 1.0 muF/cm(2) in summer and 2.0 muF/cm(2) in winter. The internal resistivity and specific membrane resistance were 167 Omega.cm and 8.9 kOmega.cm(2) in summer, and 232 Omega.cm and 3.9 kOmega.cm(2) in winter.4. Early after-depolarizations were recorded in glycerol-treated fibres which had a low membrane capacitance, did not twitch and showed little ;creep'. Electron micrographs of glycerol-treated fibres showed disruption of the transverse tubular system and sarcoplasmic reticulum.5. After exposure of muscles to 400 mM urea or acetamide for 1 hr, muscle fibres did not twitch and had a reduced membrane capacitance in Ringer solution.

Acetamides↗
PubMed1998

Linking winter and summer events in a migratory bird by using stable-carbon isotopes

For migratory birds, early arrival and physical condition on the breeding grounds are important determinants of reproductive success and fitness. Differences in arrival times often exceed a month, and later arriving individuals are often in poorer condition. Habitat-specific isotopic signatures indicate that the quality of winter habitats occupied by American redstarts (Setophaga ruticilla) determines their physical condition and spring departure dates, which in turn result in variable arrival schedules and condition on temperate breeding grounds. These findings link events in tropical winter grounds with those in temperate breeding areas for a migratory songbird and provide evidence that winter habitats may be limiting.

Journal Article↗