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Photoperiodic Regulation of the Seasonal Pattern of Photosynthetic Capacity and the Implications for Carbon Cycling
Although temperature is an important driver of seasonal changes in photosynthetic physiology, photoperiod also regulates leaf activity. Climate change will extend growing seasons if temperature cues predominate, but photoperiod-controlled species will show limited responsiveness to warming. We show that photoperiod explains more seasonal variation in photosynthetic activity across 23 tree species than temperature. Although leaves remain green, photosynthetic capacity peaks just after summer solstice and declines with decreasing photoperiod, before air temperatures peak. In support of these findings, saplings grown at constant temperature but exposed to an extended photoperiod maintained high photosynthetic capacity, but photosynthetic activity declined in saplings experiencing a naturally shortening photoperiod; leaves remained equally green in both treatments. Incorporating a photoperiodic correction of photosynthetic physiology into a global-scale terrestrial carbon-cycle model significantly improves predictions of seasonal atmospheric CO{sub 2} cycling, demonstrating the benefit of such a function in coupled climate system models. Accounting for photoperiod-induced seasonality in photosynthetic parameters reduces modeled global gross primary production 2.5% ({approx}4 PgC y{sup -1}), resulting in a >3% ({approx}2 PgC y{sup -1}) decrease of net primary production. Such a correction is also needed in models estimating current carbon uptake based on remotely sensed greenness. Photoperiod-associated declines in photosynthetic capacity could limit autumn carbon gain in forests, even if warming delays leaf senescence.
Transfer from long to short photoperiods affects production efficiency of day-neutral rice
The day-neutral, semidwarf rice (Oryza sativa L.) cultivar Ai-Nan-Tsao was grown in a greenhouse under summer conditions using high-pressure sodium lamps to extend the natural photoperiod. After allowing 2 weeks for germination, stand establishment, and thinning to a consistent planting density of 212 plants/m2, stands were maintained under continuous lighting for 35 or 49 days before shifting to 8- or 12-h photoperiods until harvest 76 days after planting. Non-shifted control treatments consisting of 8-, 12-, or 24-h photoperiods also were maintained throughout production. Tiller number increased as duration of exposure to continuous light increased before shifting to shorter photoperiods. However, shoot harvest index and yield efficiency rate were lower for all plants receiving continuous light than for those under the 8- or 12-h photoperiods. Stands receiving 12-h photoperiods throughout production had the highest grain yield per plant and equaled the 8-h-photoperiod control plants for the lowest tiller number per plant. As long as stands were exposed to continuous light, tiller formation continued. Shifting to shorter photoperiods late in the cropping cycle resulted in newly formed tillers that were either sterile or unable to mature grain before harvest. Late-forming tillers also suppressed yield of grain in early-forming tillers, presumably by competing for photosynthate or for remobilized assimilate during senescence. Stands receiving 12-h photoperiods throughout production not only produced the highest grain yield at harvest but had the highest shoot harvest index, which is important for resource-recovery strategies in advanced life-support systems proposed for space.
Identifying yield-optimizing environments for two cowpea breeding lines by manipulating photoperiod and harvest scenario
Photoperiod and harvest scenario of cowpea (Vigna unguiculata L. Walp) canopies were manipulated to optimize productivity for use in future controlled ecological life-support systems. Productivity was measured by edible yield rate (EYR:g m-2 day-1), shoot harvest index (SHI: g edible biomass [g total shoot dry weight]), and yield-efficiency rate (YER:g edible biomass m-2 day-1 per[g nonedible shoot dry weight]). Breeding lines 'IT84S-2246' (S-2246) and "IT82D-889' (D-889) were grown in a greenhouse under 8-, 12-, or 24-h photoperiods. S-2246 was short-day and D-889 was day-neutral for flowering. Under each photoperiod, cowpeas were harvested either for leaves only, seeds only, or leaves plus seeds (mixed harvest). Photoperiod did not affect EYR of either breeding line for any harvest scenario tested. Averaged over both breeding lines, seed harvest gave the highest EYR at 6.7 g m-2 day-1. The highest SHI (65%) and YER (94 mg m-2 day-1 g-1) were achieved for leaf-only harvest of D-889 under an 8-h photoperiod. For leaf-only harvest of S-2246, both SHI and YER increased with increasing photoperiod, but declined for seed-only and mixed harvests. However, photoperiod had no effect on SHI or YER for D-889 for any harvest scenario. A second experiment utilized the short-day cowpea breeding line 'IT89KD-288' (D-288) and the day-neutral breeding line 'IT87D-941-1' (D-941) to compare yield parameters using photoperiod extension under differing lamp types. This experiment confirmed the photoperiod responses of D-889 and S-2246 to a mixed-harvest scenario and indicated that daylength extension with higher irradiance from high pressure sodium lamps further suppressed EYR, SHI, and YER of the short-day breeding line D-288.
Photoperiod decelerates the advance of spring phenology of six deciduous tree species under climate warming
Abstract Vegetation phenology in spring has substantially advanced under climate warming, consequently shifting the seasonality of ecosystem process and altering biosphere–atmosphere feedbacks. However, whether and to what extent photoperiod (i.e., daylength) affects the phenological advancement is unclear, leading to large uncertainties in projecting future phenological changes. Here we examined the photoperiod effect on spring phenology at a regional scale using in situ observation of six deciduous tree species from the Pan European Phenological Network during 1980–2016. We disentangled the photoperiod effect from the temperature effect (i.e., forcing and chilling) by utilizing the unique topography of the northern Alps of Europe (i.e., varying daylength but uniform temperature distribution across latitudes) and examining phenological changes across latitudes. We found prominent photoperiod‐induced shifts in spring leaf‐out across latitudes (up to 1.7 days per latitudinal degree). Photoperiod regulates spring phenology by delaying early leaf‐out and advancing late leaf‐out caused by temperature variations. Based on these findings, we proposed two phenological models that consider the photoperiod effect through different mechanisms and compared them with a chilling model. We found that photoperiod regulation would slow down the advance in spring leaf‐out under projected climate warming and thus mitigate the increasing frost risk in spring that deciduous forests will face in the future. Our findings identify photoperiod as a critical but understudied factor influencing spring phenology, suggesting that the responses of terrestrial ecosystem processes to climate warming are likely to be overestimated without adequately considering the photoperiod effect.
Photoperiod Decelerates the Advance of Spring Phenology of Six Deciduous Tree Species under Climate Warming
Vegetation phenology in spring has substantially advanced under climate warming, consequently shifting the seasonality of ecosystem process and altering biosphere-atmosphere feedbacks. However, whether and to what extent photoperiod (i.e., daylength) affects the phenological advancement is unclear, leading to large uncertainties in projecting future phenological changes. Here we examined the photoperiod effect on spring phenology at a regional scale using in situ observation of six deciduous tree species from the Pan European Phenological Network during 1980-2016. We disentangled the photoperiod effect from the temperature effect (i.e., forcing and chilling) by utilizing the unique topography of the northern Alps of Europe (i.e., varying daylength but uniform temperature distribution across latitudes) and examining phenological changes across latitudes. We found prominent photoperiod-induced shifts in spring leaf-out across latitudes (up to 1.7 daysper latitudinal degree). Photoperiod regulates spring phenology by delaying early leaf-out and advancing late leaf-out caused by temperature variations. Based on these findings, we proposed two phenological models that consider the photoperiod effect through different mechanisms and compared them with a chilling model. We found that photoperiod regulation would slow down the advance in spring leaf-out under projected climate warming and thus mitigate the increasing frost risk in spring that deciduous forests will face in the future. Our findings identify photoperiod as a critical but understudied factor influencing spring phenology, suggesting that the responses of terrestrial ecosystem processes to climate warming are likely to be overestimated without adequately considering the photoperiod effect.
Changes in photoperiod alter Glut4 expression in skeletal muscle of C57BL/6J mice
Seasonal changes in photoperiod influence body weight and metabolism in mice. Here, we examined the effect of changes in photoperiod on the expression of glucose transporter genes in the skeletal muscle and adipose tissue of C57BL/6J mice. Glut4 expression was lower in the gastrocnemius muscle of mice exposed to a short-duration day (SD) than those to a long-duration day (LD), with accompanying changes in GLUT4 protein levels. Although Glut4 expression in the mouse soleus muscle was higher under SD than under LD, GLUT4 protein levels remained unchanged. To confirm the functional significance of photoperiod-induced changes in Glut4 expression, we checked for variations in insulin sensitivity. Blood glucose levels after insulin injection remained high under SD, suggesting that the mice exposed to SD showed lower sensitivity to insulin than those exposed to LD. We also attempted to clarify the relationship between Glut4 expression and physical activity in the mice following changes in photoperiod. Locomotor activity, as detected via infrared beam sensor, was lower under SD than under LD. However, when we facilitated voluntary activity by using running wheels, the rotation of wheels was similar for both groups of mice. Although physical activity levels were enhanced due to running wheels, Glut4 expression in the gastrocnemius muscle remained unchanged. Thus, variations in photoperiod altered Glut4 expression in the mouse skeletal muscle, with subsequent changes in GLUT4 protein levels and insulin sensitivity; these effects might be independent of physical activity. - Highlights: • Glut4 expression in the gastrocnemius muscle was lowered under short photoperiod. • Insulin sensitivity was lowered under short photoperiod. • Access to running wheels did not alter Glut4 expression in the gastrocnemius muscle. • Photoperiodic changes in Glut4 expression may be independent of physical activity.
Complex interactions between day length and diurnal patterns of gene expression drive photoperiodic responses in a perennial C 4 grass
Abstract Photoperiod is a key environmental cue affecting flowering and biomass traits in plants. Key components of the photoperiodic flowering pathway have been identified in many species, but surprisingly few studies have globally examined the diurnal rhythm of gene expression with changes in day length. Using a cost‐effective 3′‐Tag RNA sequencing strategy, we characterize 9,010 photoperiod responsive genes with strict statistical testing across a diurnal time series in the C 4 perennial grass, Panicum hallii . We show that the vast majority of photoperiod responses are driven by complex interactions between day length and sampling periods. A fine‐scale contrast analysis at each sampling time revealed a detailed picture of the temporal reprogramming of cis ‐regulatory elements and biological processes under short‐ and long‐day conditions. Phase shift analysis reveals quantitative variation among genes with photoperiod‐dependent diurnal patterns. In addition, we identify three photoperiod enriched transcription factor families with key genes involved in photoperiod flowering regulatory networks. Finally, coexpression networks analysis of GIGANTEA homolog predicted 1,668 potential coincidence partners, including five well‐known GI‐interacting proteins. Our results not only provide a resource for understanding the mechanisms of photoperiod regulation in perennial grasses but also lay a foundation to increase biomass yield in biofuel crops.
Utilization of potatoes for life support systems. II - The effects of temperature under 24-h and 12-h photoperiods
The effects of temperature and the photoperiod length on the growth and tuberization of Norland potatoes were investigated for two photoperiods, 12-h and 24-hr at 400 micromol/sq m per sec PPF, and at temperatures of 12, 16, 20, 24, and 28 C. It was found that stem length increased with increasing temperature under both photoperiods. The highest tuber yield was obtained at 16 C under the 24-hr photoperiod and at 20 C under the 12-hr photoperiod (i.e., increasing the photoperiod from 12 to 24 hrs effectively decreases the optimal temperature for tuber formation). Little or no tuber formation occurred at 28 C under either photoperiod.
Discussion of Yellow Starthistle Response to Irradiance, Photoperiod, and CO2
Yellow Starthistle (Centaurea solstitialis) is a native annual weed of Eurasia and since introduction into the US has become an invasive and noxious weed. It grows in a rosette habit during the vegetative state and usually bolts in summer to produce a large and branched flowering stem. Time to flowering in Yellow Starthistle has been attributed to photoperiod, nitrogen nutrition, temperature, and water stress. We executed a series of studies to investigate the role of light, both photoperiod and photosynthetic photon flux, on flowering and development in Yellow Starthistle. Treatments were presented in 4 ways: (1) Varying day length with constant photosynthetic photon flus (PPF) - providing increasing daily integrated Photosynthetic Photon (PP) exposure with longer day lengths, (2) Varying day length while adjusting PPF to maintain daily PP exposure for all treatments, (3) Extending photoperiod treatments beyond common 12-h photosynthetic period with low light levels to maintain both PPF and daily PP across all treatments; and (4) Reciprocal exchange of plant among photoperiod treatments. Yellow Starthistle appears to be a long-day plant with a critical day length requirement between 14-h and 16-h to induce transition from vegetative to floral stages in development. PPF and daily absorbed photons did not affect time to vegetative / floral stage transition, but did affect factors such as biomass accumulation and canopy parameters such as specific leaf mass. Reciprocal exchange of plants between floral inducing and inhibiting photoperiod treatments, starting at 2-weeks post germination, had no effect on to flower. Flowering was determined by photoperiod experienced during the first 2-weeks (or less) post germination. Yellow Starthistle net photosynthetic response to elevated atmospheric CO2 concentrations over a range of photosynthetically active radiation flux rates and temperatures will also be presented and discussed.
Discussion of Yellow Starthistle Response to Photosynthetic Irradiance, Photoperiod, and CO2
Yellow Starthistle (Centaurea solstitialis) is a native annual weed of Eurasia and since introduction into the United STates has become an invasive and noxious weed. It grows in a rosette habit during the vegetative state and usually bolts in summer to produce a large and branched flowering stem. Time to flowering in Yellow Starthistle has been attributed to photoperiod, nitrogen nutrition, temperature, and water stress. We executed a series of studies to investigate the role of light, both photoperiod and photosynthetic photon flux, on flowering and development in Yellow Starthistle. Treatments were presented in 4 ways: 1) varying day length with constant photosynthetic photon flus (PPF) providing increasing daily integrated Photosynthetic Photon (PP) exposure with longer day lengths 2) varying day length while adjusting PPF to maintain daily PP exposure for all treatments 3) extending photoperiod treatments beyond common 12-h photosynthetic period with low light levels to maintain both PPF and daily PP across all treatments4)reciprocal exchange of plant among photoperiod treatments Yellow Starthistle appears to be a long-day plant with a critical day length requirement between 14-h and 16-h to induce transition from vegetative to floral stages in development. PPF and daily absorbed photons did not affect time to vegetative floral stage transition, but did affect factors such as biomass accumulation and canopy parameters such as specific leaf mass. Reciprocal exchange of plants between floral inducing and inhibiting photoperiod treatments, starting at 2-weeks post germination, had no effect on to flower. Flowering was determined by photoperiod experienced during the first 2-weeks (or less) post germination.Yellow Starthistle net photosynthetic response to elevated atmospheric CO2 concentrations over a range of photosynthetically active radiation flux rates and temperatures will also be presented and discussed.
Temperature and photoperiod responses of soybean embryos cultured in vitro
Temperature and photoperiod each have direct effects on growth rate of excised embryos of soybean (Glycine max (L.) Merrill). To determine if the effects of photoperiod are altered by temperature, embryos of 'Ransom II' were cultured in vitro at 18, 24, and 30 degrees C under photoperiod durations of 12 and 18 h at an irradiance of 9 W m-2 (700 to 850 nm) and a photosynthetic photon flux density of 58 micromoles m-2 s-1 (400 to 700 nm). Accumulation rates of fresh and dry weight were greater under 18-h than 12-h photoperiods over the entire range of temperature. Water content of the culture embryos was not affected by photoperiod but was greater at 18 and 30 than 24 degrees C. The accumulation rate of dry weight increased from 18 to 26 but declined at 30 degrees C.
Internode and petiole elongation of soybean in response to photoperiod and end-of-day light quality
Elongation of main stem internodes and petioles of soybeans, Glycine max 'Ransom,' was examined in response to various photoperiod/temperature combinations and to end-of-day (EOD) light quality. Photoperiod treatments consisted of 10, 14, and 16 h in combination with day/night temperatures of 18/14, 22/18, 26/22, 30/26, and 34/30 C. The EOD treatments consisted of exposing plants to illumination from either incandescent (high far-red component, FR) or fluorescent (high red component, R) lamps during the final 0.5 h of a 10-h photoperiod. Internode elongation was not significantly promoted by the photoperiod treatments, and, in fact, under the two highest temperature regimes, internode elongation was suppressed under the longer photoperiods. Petiole elongation, however, was enhanced under the longer photoperiods at all temperatures. In the EOD light study, internode and petiole elongation was significantly greater on plants exposed to 0.5 h EOD from incandescent lamps than from fluorescent. Under the incandescent EOD treatment, plants increased dry matter production by 41% and exhibited greater partitioning of assimilates in stem and root portions than under fluorescent EOD.
EARLY FLOWERING 3 and Photoperiod Sensing in Brachypodium distachyon
The proper timing of flowering, which is key to maximize reproductive success and yield, relies in many plant species on the coordination between environmental cues and endogenous developmental programs. The perception of changes in day length is one of the most reliable cues of seasonal change, and this involves the interplay between the sensing of light signals and the circadian clock. Here, we describe a Brachypodium distachyon mutant allele of the evening complex protein EARLY FLOWERING 3 (ELF3). We show that the elf3 mutant flowers more rapidly than wild type plants in short days as well as under longer photoperiods but, in very long (20 h) days, flowering is equally rapid in elf3 and wild type. Furthermore, flowering in the elf3 mutant is still sensitive to vernalization, but not to ambient temperature changes. Molecular analyses revealed that the expression of a short-day marker gene is suppressed in elf3 grown in short days, and the expression patterns of clock genes and flowering time regulators are altered. We also explored the mechanisms of photoperiodic perception in temperate grasses by exposing B. distachyon plants grown under a 12 h photoperiod to a daily night break consisting of a mixture of red and far-red light. We showed that 2 h breaks are sufficient to accelerate flowering in B. distachyon under non-inductive photoperiods and that this acceleration of flowering is mediated by red light. Finally, we discuss advances and perspectives for research on the perception of photoperiod in temperate grasses.
Effects of CO2 Concentration on Leaf Photosynthesis and Stomatal Conductance of Potatoes Grown Under Different Irradiance Levels and Photoperiods
Potato (Solanum tuberosum L.) cvs. Russet Burbank, Denali, and Norland, were grown in environmental rooms controlled at approx 350 micro mol/mol (ambient during years 1987/1988) and 1000 micro mol/mol (enriched) CO2 concentrations. Plants and electric lamps were arranged to provide two irradiance zones, 400 and 800 micro mol/mol/square m/S PPF and studies were repeated using two photoperiods (12-h light / 12-h dark and continuous light). Leaf photosynthetic rates and leaf stomatal conductance were measured using fully expanded, upper canopy leaves at weekly intervals throughout growth (21 through 84 days after transplanting). Increasing the CO2 from approx 350 to 1000 micro mol/mol under the 12-h photoperiod increased leaf photosynthetic rates by 39% at 400 micro mol/mol/square m/S PPF and 27% at 800 micro mol/mol/square m/S PPF. Increasing the CO2 from approx 350 to 1000 micro mol/mol under continuous light decreased leaf photosynthetic rates by 7% at 400 micro mol/mol/square m/S PPF and 13% at 800 micro mol/mol/square m/S PPF. Increasing the CO2 from approx 350 to 1000 micro mol/mol under the 12-h photoperiod plants decreased stomatal conductance by an average of 26% at 400 micro mol/mol/square m/S PPF and 42% at 800 micro mol/mol/square m/S PPF. Under continuous light, CO2 enrichment resulted in a small increase (2%) of stomatal conductance at 400 micro mol/mol/square m/S PPF, and a small decrease (3%) at 800 micro mol/mol/square m/S PPF. Results indicate that CO2 enrichment under the 12-h photoperiod showed the expected increase in photosynthesis and decrease in stomatal conductance for a C3 species like potato, but the decreases in leaf photosynthetic rates and minimal effect on conductance from CO2 enrichment under continuous light were not expected. The plant leaves under continuous light showed more chlorosis and some rusty flecking versus plants under the 12-h photoperiod, suggesting the continuous light was more stressful on the plants. The increased rates of leaf photosynthesis with increased CO2 concentration paralleled trends in biomass production (published previously) but were not proportional to the biomass yields.
Interacting effects of photoperiod and photosynthetic photon flux on net carbon assimilation and starch accumulation in potato leaves
The effect of photoperiod (PP) on net carbon assimilation rate (Anet) and starch accumulation in newly mature canopy leaves of 'Norland' potato (Solanum tuberosum L.) was determined under high (412 varies as mol m-2s-1) and low (263 varies as mol m-2s-1) photosynthetic photon flux (PPF) conditions. The Anet decreased from 13.9 to 11.6 and 9.3 micromoles m-2s-1, and leaf starch increased from 70 to 129 and 118 mg g-1 drymass (DM) as photoperiod (PP) was increased from 12/12 to 18/6, and 24/0, respectively. Longer PP had a greater effect with high PPF conditions than with low PPF treatments, with high PPF showing greater decline in Anet. Photoperiod did not affect either the CO2 compensation point (50 micromoles mol-1) or CO2 saturation point (1100-1200 micromoles mol-1) for Anet. These results show an apparent limit to the amount of starch that can be stored (approximately 15% DM) in potato leaves. An apparent feedback mechanism exists for regulating Anet under high PPF, high CO2, and long PP, but there was no correlation between Anet and starch concentration in individual leaves. This suggests that maximum Anet cannot be sustained with elevated CO2 conditions under long PP (> or = 12 hours) and high PPF conditions. If a physiological limit exists for the fixation and transport of carbon,then increasing photoperiod and light intensity under high CO2 conditions is not the most appropriate means to maximize the yield of potatoes.
Prolonging photoperiod promotes testosterone synthesis of Leydig cells by directly targeting local melatonin system in rooster testes
Abstract The study investigated the effects of prolonging photoperiod on the synthesis of testosterone and melatonin in roosters, and the effect of melatonin on testosterone synthesis in rooster Leydig cells as well as its molecular mechanisms. We randomly divided one hundred and twenty 20-week-old roosters into three groups and provided 6, 12.5 and 16 h light, respectively. The results showed that prolonging photoperiod promoted testosterone synthesis, decreased melatonin production, and inhibited the expression of melatonin membrane receptors MEL1A, MEL1B, MEL1C, and aralkylamine n-acetyltransferase (AANAT) in rooster testes. Subsequently, rooster Leydig cells were isolated and treated with 0, 0.1, 1, 10, and 100 ng/mL melatonin for 36 h. The results suggested that melatonin inhibited testosterone synthesis in rooster Leydig cells, and silencing MEL1A and MEL1B relieved the inhibition of melatonin on testosterone synthesis. Additionally, melatonin reduced the intracellular cyclic adenosine monophosphate (cAMP) level and the phosphorylation level of cAMP-response element binding protein (CREB), and CREB overexpression alleviated the inhibition of melatonin on testosterone synthesis. Furthermore, pretreatment with cAMP activator forskolin or protein kinase A (PKA) activator 8-bromo-cAMP blocked the inhibition of melatonin on CREB phosphorylation and testosterone synthesis. These results indicated that prolonging photoperiod promoted testosterone synthesis associated with the decrease in melatonin production and membrane receptors and biosynthetic enzyme of melatonin in rooster testes, and melatonin inhibited testosterone synthesis of rooster Leydig cells by inhibiting the cAMP/PKA/CREB pathway via MEL1A and MEL1B. This may be evidence that prolonging photoperiod could promote testosterone synthesis through the inhibition of the local melatonin pathway in rooster testes.
Cold acclimation threshold induction temperatures of switchgrass ecotypes grown under a long and short photoperiod
Abstract Plants can cold acclimate to enhance their freezing tolerance by sensing declining temperature and photoperiod cues. However, the factors influencing genotypic variation in the induction of cold acclimation are poorly understood among perennial grasses. We hypothesized that the more northern upland switchgrass ( Panicum virgatum L.) ecotype develops a higher degree of freezing tolerance by initiating cold acclimation at higher temperatures as compared with the coastal and southern lowland ecotypes. First, we determined the optimal method for assessing freezing tolerance and the length of exposure to 8/4°C required to induce the maximum level of freezing tolerance in the most northern upland and most southern lowland genotypes. We characterized the maximum freezing tolerance of eight uplands, three coastal and five lowland genotypes grown for 21 days at 8/4°C and a 10 or 16 h photoperiod. Next, we identified the temperature required to induce cold acclimation by exposing the 16 genotypes for 7 days at 20–6°C constant temperatures under a 10 or 16 h photoperiod. Cold acclimation initiated at temperatures 5 and 7°C higher in upland than in coastal and lowland genotypes. Among upland genotypes the shorter photoperiod induced cold acclimation at a 1°C higher temperature. Genotypes originating from a more northern latitude initiate cold acclimation at higher temperatures and develop higher maximum freezing tolerances. An earlier response to declining temperatures may provide the upland ecotype with additional time to prepare for winter and provide an advantage when plants are subjected to the rapid changes in fall temperature associated with injurious frosts.