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Winter profile of plasma sex steroid levels in free-living male western diamond-backed rattlesnakes, Crotalus atrox (Serpentes: Viperidae).

Recent field studies on the reproductive ecology of western diamond-backed rattlesnakes (Crotalus atrox) from populations in southern Arizona showed significant differences in the concentration of plasma sex steroids (testosterone, T; 5alpha-dihydrotestosterone, DHT; and 17beta-estradiol, E2) throughout the active season (March-October), and peak levels were coincident with the two mating periods (late summer and early spring). There is, however, no information on levels of sex steroids during winter. Similar to most snakes, hibernating individuals of C. atrox are typically inaccessible, but in southern Arizona, where environmental conditions are typically mild during winter, adult males frequently bask at or near the entrances of communal dens. Basking activity, therefore, offers a unique logistical opportunity to assess the complete annual profile of plasma sex steroid levels in males of a temperate reptile in nature. From November to February, we measured levels of plasma T, DHT, and E2 in adult male C. atrox that were located basking at communal dens. Additionally, cloacal, core body, and ambient air temperatures were obtained to investigate potential relationships between body temperatures and levels of sex steroids. Mean levels of T, DHT, and E2 were relatively high, and the concentration hierarchy was T>DHT>E2. Mean levels of T, DHT, and E2 showed no significant variation across the four months of sampling; however, E2 levels decreased progressively. In the annul cycle, sex steroid levels during winter were not basal when compared to values obtained during the active season. Mean cloacal temperatures of basking males were significantly higher than core body temperatures of non-basking males (inside dens) from November-December, and in February, which suggests that one function of winter basking is to elevate body temperatures. Steroid levels, nonetheless, were not significantly correlated with cloacal temperatures. We suggest that future field studies of male C. atrox should: (a) investigate sex steroid levels in non-basking individuals and (b) test whether elevated levels of sex steroids during winter facilitate the large increases that occur in early spring, which are coincident with the second mating season. Our findings on the reproductive biology of C. atrox and other viperids are discussed in the context of the associated-dissociated model of reproduction.

Animals↗

Evidence for continued transmission of parasitic nematodes in reindeer during the Arctic winter.

Living in the high Arctic, the Svalbard reindeer (Rangifer tarandus platyrhynchus) and its trichostrongyle nematodes experience a long cold winter from October to late May/early June. Over this period, transmission would be expected to be low. However, in culled reindeer the abundance of infection increased from autumn to late winter, providing evidence for continued transmission within this period. To our knowledge this is the first time this has been demonstrated in a climate with temperatures consistently below 0 degrees C. In one winter (1996-1997), the average fraction of nematodes found as larvae in the abomasal mucosa increased from around 10% to 50% between October and March. This suggests that arrested development took place throughout the winter. We found no evidence for an efficient acquired immune response towards the nematodes. The abundance of infection did not tend to decrease with increasing host age after an earlier peak, but levelled off instead, as predicted by a simple immigration-death model. In the late winter when the nutritional plane is low, both adult reindeer and calves had high worm burdens at intensities that may affect their condition and fitness.

Abomasum↗

Low total body bone mineral content and high bone resorption in Korean winter-born versus summer-born newborn infants.

Seasonal differences in newborn total body bone mineral content (TBBMC) have not been studied, particularly in relation to alterations in vitamin D status in winter. In vitamin D deficiency bone resorption may be high and bone mineralization low. Bone resorption may be assessed by serum cross-linked carboxyterminal telopeptide of type I collagen (ICTP) measures. Because vitamin D supplements throughout pregnancy are uncommon in Korea, we hypothesized that in Korean winter newborns, TBBMC is low and serum ICTP high from high bone resorption and low 25-hydroxyvitamin D (25-OHD) compared with those in summer newborns. Seventy-one Korean term infants were studied prospectively in summer (July through September, n = 37) versus winter (January through March, n = 34); TBBMC was measured before 3 days of age by dual-energy x-ray absorptiometry. Significant seasonal differences were found: winter newborns had 6% lower TBBMC (least squares means +/- SD; 86.7 +/- 7.7 gm vs 93.9 +/- 7.8 gm, p = 0.0002), lower cord serum 25-OHD (10.7 +/- 8 nm vs 30 +/- 15 nm, p = 0.0001) and 1,25-dihydroxyvitamin D, and higher ICTP (96.4 +/- 20.3 microg/L vs 74.8 +/- 24 microg/L, p = 0.0002) and calcium than summer newborns. TBBMC correlated with serum 25-OHD (r = 0.243, p = 0.047) and inversely with ICTP (r = -0.333, p = 0.008). We suggest that in Korea low maternal vitamin D status in winter results in marked reduction in newborn TBBMC.

Absorptiometry, Photon↗

Seasonal variations of plasma fibrinogen and factor VII activity in the elderly: winter infections and death from cardiovascular disease.

There are approximately 20,000 excess deaths from cardiovascular disease each winter in England and Wales. The reasons for the excess have not been fully elucidated. For one year, we studied 96 men and women aged 65-74 living in their own homes in order to examine seasonal variation in plasma fibrinogen and factor VII clotting activity (FVIIc), and to investigate relationships with infection and other cardiovascular-disease risk factors. Both fibrinogen and FVIIc plasma values were greater in winter with estimated winter-summer differences (confidence intervals) of 0.13 (0.05-0.20) g/L for fibrinogen and 4.2 (1.2-7.1)% of standard for FVIIc. These differences could account for 15% and 9% increases in ischaemic heart disease risk in winter respectively. After adjustment for confounding by season, fibrinogen was strongly related to neutrophil count (p < 0.0001), C-reactive protein (p < 0.0001), alpha 1-antichymotrypsin (p < 0.0001), and self-reported cough (p < 0.0001) and coryza (p = 0.0004), but not to ambient temperature. Therefore, we suggest that seasonal variation in fibrinogen might be induced by winter respiratory infections via activation of the acute phase response. Seasonal variations in the cardiovascular risk factors fibrinogen and FVIIc provide further possible explanations for the marked seasonal variation in death from ischaemic heart disease and stroke in the elderly.

Acute-Phase Proteins↗

Electrophysiological evidence suggesting a seasonal modulation of retinal sensitivity in subsyndromal winter depression.

BACKGROUND: An anomaly in the retinal adaptation processes to the decreased light exposure in winter has been suggested as a contributing factor in winter depression. The purpose of this study was to investigate seasonal variations in rod sensitivity in normal subjects and in subjects with seasonal mood variations. METHODS: Nine normal subjects (5 men, 4 women, aged 21-28 years) and 12 subjects with subsyndromal seasonal affective disorder (S-SAD)(3 men, 9 women, aged 21-44 years) were selected based on their global seasonality score (GSS) from the Seasonal Pattern Assessment Questionnaire. Scotopic electroretinograms (ERGs) were obtained once in winter and once in summer. Retinal sensitivity, which represents a relative threshold, was obtained from the rod ERG luminance-response functions. RESULTS: A difference in retinal sensitivity between the two groups appeared only in the winter with lower retinal sensitivity found in the S-SAD group. A positive correlation between the GSS and the magnitude of the winter decrease in rod sensitivity was also observed. LIMITATIONS: The S-SAD subjects studied in this research did not receive a formal psychiatric evaluation. This will be necessary in future studies to determine if the changes in retinal sensitivity are specific to seasonal affective disorders. In addition, in the present study, the differences in age and gender between the two groups limit the interpretation of the possible contribution of these two parameters to the results. CONCLUSION: The seasonal changes in retinal sensitivity that parallel seasonal mood variations suggest that the ERG may represent a useful tool to investigate seasonal affective disorders.

Adaptation, Ocular↗

Energy level in winter diets of Fallow deer: effect on plasma levels of insulin-like growth factor-I and sex ratio of their offspring.

To determine the effect of dietary energy level in the first winter on subsequent puberty onset, pregnancy, growth, and secretion of progesterone and insulin-like growth factor-I (IGF-I) in Fallow deer, prepubertal deer were fed either a high (H) or a low (L) energy diet in a randomised complete block design. June-born female Fallow deer were fed ad libitum either 12.5MJ/kg DM (H, n=29) or 10MJ/kg DM (L, n=29) in pelleted rations once per day during the winter (approximately 4-10 months of age) preceding puberty. Blood samples were collected twice weekly during the peripubertal period. During the winter feeding period, DM intake was similar for both groups but average daily gain was greater (P<0.05) for deer fed the H versus L diet. Onset of puberty was not affected (P>0.10) by dietary treatment. Concentrations of progesterone in plasma did not differ (P>0.10) between dietary treatments before or after puberty, increasing after puberty in both groups, and reaching maximal levels 8-12 days after the onset of puberty. Concentrations of plasma IGF-I increased (P<0.05) before puberty in both groups reaching maximal levels 3-4 days before the onset of puberty but did not differ (P>0.10) between H and L diets before or after puberty. Of the 28 does fed the H diet that calved, 75% of the calves born were male versus 46% in the L diet (P<0.05). In conclusion, increased plasma IGF-I concentrations were associated with the onset of puberty in Fallow deer regardless of the level of dietary energy intake during the preceding winter. Increased dietary energy during winter does not alter pregnancy rates but does alter sex ratio of calves born.

Journal Article↗

Serum prolactin and dehydroepiandrosterone concentrations during the summer and winter hair growth cycles of mink (Mustela vison).

We investigated the relationship between serum concentrations of prolactin (PRL) and dehydroepiandrosterone (DHEA) during initiation and development of summer and winter hair growth (anagen) cycles in mink. In the spring, haloperidol (HAL) increased PRL concentrations and induced summer anagen earlier than controls, whereas melatonin (MEL) inhibited PRL secretion and completely blocked summer anagen. In the fall, HAL increased PRL concentrations, inducing anagen at an earlier time than controls, although the resulting fur was abnormal being almost devoid of underhair fibers. Exogenous MEL during the fall reduced PRL concentrations, initiating winter anagen 4 weeks earlier than controls. Adrenalectomy (ADX) induced earlier onset of summer and winter anagen and neutralized the inhibitory effects of HAL in the fall and MEL in the spring. No change in serum DHEA concentrations was observed during the onset of summer or winter anagen in any group although MEL increased DHEA levels from 27 March through 5 June relative to HAL-treated mink. We conclude that changes in serum levels of DHEA and PRL are not requisite to onset of summer or winter anagen in mink. It is possible that metabolites of DHEA and/or PRL may still affect other aspects of the hair growth cycle.

Adrenal Glands↗

Bone metabolic changes in Antarctic wintering team members.

Bone mass, bone metabolic markers, and calcium regulation hormones were measured in members of an Antarctic wintering team who stayed at the Japanese Antarctic station, Syowa (latitude: south 69 degrees 00', longitude: east 39 degrees 35') for 1 year. Subjects included 31 healthy Japanese males, aged 24-51 years (mean age 34.5 years) at the beginning of this study, ingesting 488 IU/day of vitamin D and 550.9 mg/day of calcium per person. The long-term coefficient of variation (CV) of the equipment used in the assessments of bone mass was 0.67% in single X-ray absorptiometry (SXA), 0.17% in the speed of sound (SOS) by quantitative ultrasound method (QUS), and 0.63% in broadband ultrasound attenuation (BUA) by QUS. The seasonal changes in the calcaneal bone mineral density (BMD) by SXA were not significant, whereas the SOS measured by QUS decreased during the measurement period (0.55%, p < 0.001), and BUA increased (1.9%, p < 0.01). Bone-specific alkaline phosphatase and osteocalcin levels increased significantly during summer (p < 0.001) and urinary calcium level decreased significantly during winter (p < 0.05). Urinary pyridinoline and deoxypyridinoline levels decreased significantly at the end of winter (p < 0.001). Serum 1,25(OH)2D3 level did not change significantly, whereas serum 25(OH)D3 level decreased significantly during winter (p < 0.001). Serum parathyroid hormone (PTH) level significantly increased at the end of winter (p < 0.01), although both PTH level and 25(OH)D3 level remained within the normal range. We concluded that the 25(OH)D3 level in subjects who stayed in Antarctica for 1 year decreased significantly with the reduction in duration of sunshine, but there were no clear changes in bone mass.

Adult↗

Winter forest soil respiration controlled by climate and microbial community composition.

Most terrestrial carbon sequestration at mid-latitudes in the Northern Hemisphere occurs in seasonal, montane forest ecosystems. Winter respiratory carbon dioxide losses from these ecosystems are high, and over half of the carbon assimilated by photosynthesis in the summer can be lost the following winter. The amount of winter carbon dioxide loss is potentially susceptible to changes in the depth of the snowpack; a shallower snowpack has less insulation potential, causing colder soil temperatures and potentially lower soil respiration rates. Recent climate analyses have shown widespread declines in the winter snowpack of mountain ecosystems in the western USA and Europe that are coupled to positive temperature anomalies. Here we study the effect of changes in snow cover on soil carbon cycling within the context of natural climate variation. We use a six-year record of net ecosystem carbon dioxide exchange in a subalpine forest to show that years with a reduced winter snowpack are accompanied by significantly lower rates of soil respiration. Furthermore, we show that the cause of the high sensitivity of soil respiration rate to changes in snow depth is a unique soil microbial community that exhibits exponential growth and high rates of substrate utilization at the cold temperatures that exist beneath the snow. Our observations suggest that a warmer climate may change soil carbon sequestration rates in forest ecosystems owing to changes in the depth of the insulating snow cover.

Bacteria↗

Physical and transcriptional map of the critical region for keratolytic winter erythema (KWE) on chromosome 8p22-p23 between D8S550 and D8S1759.

Keratolytic winter erythema is an autosomal dominant skin disorder characterised by erythema, hyperkeratosis, and peeling of the skin of the palms and soles, especially during winter. The keratolytic winter erythema locus has been mapped to human chromosome 8p22-p23. This chromosomal region has also been associated with frequent loss of heterozygosity in different types of cancer. To identify positional candidate genes for keratolytic winter erythema, a BAC contig located between the markers at D8S550 and D8S1695 was constructed and sequenced. It could be extended to D8S1759 by a partially sequenced BAC clone identified by database searches. In the 634 404 bp contig 13 new polymorphic microsatellite loci and 46 single nucleotide and insertion/deletion polymorphisms were identified. Twelve transcripts were identified between D8S550 and D8S1759 by exon trapping, cDNA selection, and sequence analyses. They were localised on the genomic sequence, their exon/intron structure was determined, and their expression analysed by RT-PCR. Only one of the transcripts corresponds to a known gene, encoding B-lymphocyte specific tyrosine kinase, BLK. A putative novel myotubularin-related protein gene (MTMR8), a potential human homologue of the mouse acyl-malonyl condensing enzyme gene (Amac1), and two transcripts showing similarities to the mouse L-threonine 3-dehydrogenase gene and the human SEC oncogene, respectively, were identified. The remaining seven transcripts did not show similarities to known genes. There were no potentially pathogenic mutations identified in any of these transcripts in keratolytic winter erythema patients.

Chromosomes, Artificial, Bacterial↗

Sleep stage and skin temperature regulation during night-sleep in winter.

To clarify the thermally comfortable environment for satisfied sleep, we investigated the relationship between skin temperatures and sleep stages during night-sleep in winter, summer and autumn. Four healthy young females served as the subjects. The experiments were performed in a climate chamber under three conditions: at 13 degrees C-RH70% with futon and blanket in winter, at 20-18 degrees C-RH60% with futon and towelket in autumn and at 29 degrees C-RH70% with towelket in summer. The subjects' skin temperature on the sole was lower until about 4 hours after lights off in winter than in autumn and summer, and the percentage of st.2 was lower and st.4 was higher in winter than in autumn and summer, although the bed climate both in winter and autumn was much the same.

Adult↗

Improved antioxidative protection in winter swimmers.

Adaptation to oxidative stress is an improved ability to resist the damaging effects of reactive oxygen species, resulting from pre-exposure to a lower dose. Changes in uric acid and glutathione levels during ice-bathing suggest that the intensive voluntary short-term cold exposure of winter swimming produces oxidative stress. We investigated whether the repeated oxidative stress in winter swimmers results in improved antioxidative adaptation. We obtained venous blood samples from winter swimmers and determined important components of the antioxidative defense system in the erythrocytes or blood plasma: reduced and oxidized glutathione (GSH and GSSG), and the activities of superoxide dismutase (SOD), glutathione peroxidase (GPx) and catalase (Cat). The control group consisted of healthy people who had never participated in winter swimming. The baseline concentration of GSH and the activities of erythrocytic SOD and Cat, were higher in winter swimmers. We interpret this as an adaptative response to repeated oxidative stress, and postulate it as a new basic molecular mechanism of increased tolerance to environmental stress.

Adaptation, Physiological↗

Effects of summer drought and winter freezing on stem hydraulic conductivity of Rhododendron species from contrasting climates.

We studied the limits to maximum water transport in three diffuse-porous evergreen shrubs exposed to frequent winter freeze-thaw events (Rhododendron maximum L. and R. catawbiense Michaux from the Appalachian Mountains) and to a severe summer drought (R. macrophyllum G. Don. from the Oregon Cascades). Percent loss of hydraulic conductivity (PLC), vulnerability curves to xylem embolism and freezing point temperatures of stems were measured over 2 years. Controlled freeze-thaw experiments were also conducted to determine the effect of thaw rate on PLC. During both years, native PLC was significantly higher in winter than in summer for R. macrophyllum. Seasonal changes in PLC were variable in both R. catawbiense and R. maximum. Only R. maximum plants growing in gaps or clearings showed higher PLC than understory plants. A rapid (2-4 day) natural recovery of high native PLC during the winter was observed in both R. maximum and R. macrophyllum. Compared with the bench-dehydration method, vulnerability curves based on the air-injection method consistently had less negative slopes and greater variation. Fifty percent PLC (PLC(50)) obtained from vulnerability curves based on the dehydration method occurred at -1.75, -2.42 and -2.96 MPa for R. catawbiense, R. maximum and R. macrophyllum, respectively. Among the study species, R. macrophyllum, which commonly experiences a summer drought, had the most negative water potential at PLC(50). In all species, stem freezing point temperatures were not consistently lower in winter than in summer. A single fast freeze-thaw event significantly increased PLC, and R. catawbiense had the highest PLC in response to freezing treatments. Recovery to control PLC values occurred if a low positive hydraulic pressure was maintained during thawing. Rhododendron macrophyllum plants, which commonly experience few freeze-thaw events, had large stem diameters, whereas plants of R. catawbiense, which had small stem diameters, suffered high embolism in response to a single freeze-thaw event. Both drought-induced and winter-induced embolism caused a significant reduction in hydraulic conductivity in all species during periods when drought or freeze-thaw events occurred in their native habitats. However, rapid recovery of PLC following freezing or drought maintained the species above their relatively low margins of safety for complete xylem dysfunction.

Climate↗

Winter photoinhibition in the field involves different processes in four co-occurring Mediterranean tree species.

Photoinhibition was examined in four co-occurring Mediterranean evergreen tree species during two consecutive winters. In response to low temperatures and saturating light, Juniperus phoenicea L., Pinus halepensis Mill., Quercus coccifera L. and Q. ilex ssp. ballota (Desf.) Samp. exhibited marked chronic photoinhibition, indicated by low predawn maximal photochemical efficiency of photosystem II (PSII) (Fv/Fm). Low Fv/Fm values were correlated with high concentrations of xanthophyll cycle components (VAZ) and with the maintenance of high concentrations of zeaxanthin overnight (DPSpd). In all species, however, chronic photoinhibition was enhanced as the winter progressed in the absence of changes in DPSpd, suggesting cumulative damage toward the end of winter. Photoinhibition differed among species: P. halepensis always displayed significantly higher Fv/Fm values; and Q. coccifera had the lowest Fv/Fm values, showing a high sensitivity to the combination of high light and low temperatures. Differences among species were not fully explained by differences in the xanthophyll pool or its de-epoxidation state. Chronic photoinhibition overlapped with a dynamic photoinhibition as shown by the low values of photochemical efficiency of the open reaction centers of PSII at midday. Winter photoprotective strategies differed among species and may involve photoprotective mechanisms in addition to those associated with xanthophylls. The observed species-specific differences matched results obtained for the same species in summer; however, comparison of the two seasons suggests that the higher VAZ concentration observed in winter has an additional structural photoprotective role.

Chlorophyll↗

Stable isotope evidence of diverse species-specific and individual wintering strategies in seabirds.

Although there is increasing evidence that climatic variations during the non-breeding season shape population dynamics of seabirds, most aspects of their winter distribution and ecology remain essentially unknown. We used stable isotope signatures in feathers to infer and compare the moulting (wintering) habitat of subantarctic petrels breeding at two distant localities (South Georgia and Kerguelen). Petrels showed species-specific wintering habitat preferences, with a similar pattern of latitudinal segregation for all but one taxon. At both localities, delta13C values indicated that blue petrels (Halobaena caerulea) moult in Antarctic waters, South Georgian diving petrels (Pelecanoides georgicus) in the vicinity of the archipelagos and/or in the Polar Frontal Zone and Antarctic prions (Pachyptila desolata) in warmer waters. In contrast, common diving petrels (Pelecanoides urinatrix) showed divergent strategies, with low and high intrapopulation variation at South Georgia and Kerguelen, respectively. Birds from Kerguelen dispersed over a much wider range of habitats, from coastal to oceanic waters and from Antarctica to the subtropics, whereas those from South Georgia wintered mainly in waters around the archipelago. This study is the first to show such striking between-population heterogeneity in individual wintering strategies, which could have important implications for likely demographic responses to environmental perturbation.

Animal Migration↗

Differential migration and an endocrine response to stress in wintering dark-eyed juncos (Junco hyemalis).

The dark-eyed junco (junco hyemalis) exhibits differential migration in autumn that, in general, results in females overwintering south of males, and young within each sex overwintering north of older birds. Individuals overwintering at higher latitudes face less predictable and more challenging environmental conditions. Rapid increases in circulating levels of the energy-regulating glucocorticosteroid, corticosterone, occur in response to environmental stressors. To establish whether the strength of acute corticosterone secretion was correlated with the probability of encountering poor environmental conditions, we compared the corticosterone stress response (e.g. initial plasma concentrations at the time of capture and 30 min later) in dark-eyed juncos overwintering in Mississippi (MS), USA, near the southern limit of their wintering range, with juncos overwintering in New York (NY), USA, near the northern limit of their wintering range. During two winters, 22 males and one female were sampled in NY; 13 males, 12 females and one bird of undetermined sex were sampled in MS. Not unexpectedly, NY birds carried greater fat reserves that resulted in a significantly higher value of energetic condition (mass corrected for wing cord cubed). There was no difference between the two winters sampled at either site, nor was there an effect of sex on patterns of corticosterone secretion in MS birds. With sexes pooled, MS and NY birds had similar baseline corticosterone levels. However, as predicted, NY birds exhibited significantly higher corticosterone concentrations 30 min after capture. These results support the hypothesis that birds wintering in less predictable, more extreme environments show a higher amplitude corticosterone response, which may enable them to adjust their behaviour and physiology more rapidly in response to environmental stressors such as storms. Adrenocortical sensitivity may be a part of the physiological milieu associated with differential migration in juncos; whether it results from endogenous differences in the migratory programmes of individuals or from acclimatization to local environmental conditions remains to be determined.

Adaptation, Physiological↗

Winter availability of cereal stubbles attracts declining farmland birds and positively influences breeding population trends.

Many studies have demonstrated the selection of stubble fields by farmland birds in winter, but none have shown whether provisioning of this key habitat positively influences national population trends for widespread farmland birds. We use two complementary extensive bird surveys undertaken at the same localities in summer and winter and show that the area of stubble in winter attracts increased numbers of several bird species of conservation concern. Moreover, for several farmland specialists, the availability of stubble fields in winter positively influenced the 10 year breeding population trend (1994-2003) whereas hedgerow bird species were less affected. For skylarks and yellowhammers, initially negative trends showed recovery with 10-20 ha of stubble per 1 km square. Thus, agri-environment schemes that promote retention of over-winter stubbles will attract birds locally and are capable of reversing current population declines if stubbles are available in sufficient quantity.

Agriculture↗

Integration of flowering signals in winter-annual Arabidopsis.

Photoperiod is the primary environmental factor affecting flowering time in rapid-cycling accessions of Arabidopsis (Arabidopsis thaliana). Winter-annual Arabidopsis, in contrast, have both a photoperiod and a vernalization requirement for rapid flowering. In winter annuals, high levels of the floral inhibitor FLC (FLOWERING LOCUS C) suppress flowering prior to vernalization. FLC acts to delay flowering, in part, by suppressing expression of the floral promoter SOC1 (SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1). Vernalization leads to a permanent epigenetic suppression of FLC. To investigate how winter-annual accessions integrate signals from the photoperiod and vernalization pathways, we have examined activation-tagged alleles of FT and the FT homolog, TSF (TWIN SISTER OF FT), in a winter-annual background. Activation of FT or TSF strongly suppresses the FLC-mediated late-flowering phenotype of winter annuals; however, FT and TSF overexpression does not affect FLC mRNA levels. Rather, FT and TSF bypass the block to flowering created by FLC by activating SOC1 expression. We have also found that FLC acts as a dosage-dependent inhibitor of FT expression. Thus, the integration of flowering signals from the photoperiod and vernalization pathways occurs, at least in part, through the regulation of FT, TSF, and SOC1.

Arabidopsis↗