Geographic variation as a result of evolution of the traits with broad and narrow norms of reaction in the moor frog (Rana arvalis)

Females reproductive, size, and age characteristics were studied in isolated local populations of Rana arvalis in the southern and northern parts of its range. The yearlings of the southern populations used to get larger by their first overwintering due to earlier beginning of the breeding season, as compared with the yearlings of the northern population. As a result, "southern" females become sexually mature at the age of two years while the "northern" ones become mature at the age of three years. This causes geographic differences in age composition among two populations, the "southern" reproductive females being younger on average than the "northern" ones. The earlier female maturation in the first case is not compensated by respective rise of the growth rate; to the contrary, the "southern" females grow more slowly during the first two years of their life and appear to be smaller than the "norhern" ones. These reproduction and growth patterns arise supposedly due to paedomorphosis, which causes specific reproductive characteristics, namely decrease in the egg size, increase in the reproductive effort and more strong correlation between female fertility and body size. Local and geographic differences are expressed not in the extent but in the structure of reproductive pattern, as no negative correlation was revealed between female reproductive age and body size in the southern populations. Southern habitats cannot be considered as "unfavourable with respect to body size", so the geographic differences under consideration cannot be explained by optimization of the reproductive strategies at population level. Paedomorphosis appears as a result of the female maturation rate possessing a wider norm of reaction than the growth rate. At the same time, fixation of the specific growth rate narrows norm of reaction of some other characters important for the phenotype reproductive fitness thus predetermining their subsequent evolution.     

Growth and body size in populations of mink frogs Rana septentrionalis from two latitudes

Age, longevity, and growth in mink frogs Rana septentrionalis from two latitudes in Quebec, Canada, were assessed by skeletochronological and back-calculation methods in order to document proximate causes for intraspecific variations in adult body size. In both study sites, females grew faster and were on average 11% larger than males. Mean age and maximal longevity were significantly higher in females than in males only in the southern populations. There was thus an interpopulation difference in the relative contribution of age and growth to sexual size dimorphisms. Sex-ratio also favored females (males sulfering higher mortality) only in the southern populations. Specimens from the northern population had higher mean ages (but not higher longevities) and were 17% larger than specimens from the southern populations. Annual growth rate appeared similar at the two study sites despite a shorter growing season at the northern locality. Maturity was reached by both sexes after 1 yr of post-metamorphic life (PML) in the southern populations but after 2 yr of PML in the northern population. There are indications that tadpoles metamorphose at larger body size at the northern locality after a prolonged larval period. It is concluded that growth rate, delayed maturity, greater mean ages, and eventually size at transformation, all contribute to the larger size of adult mink frogs at northern localities.     

Spatial variation in the effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus

The effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus populations were compared between coral reefs open or closed (no‐take marine reserves) to fishing and among four geographic regions of the Great Barrier Reef (GBR), Australia. The specific reproductive metrics investigated were the sex ratio, the proportion of vitellogenic females and the spawning fraction of local populations. Sex ratios became increasingly male biased with length and age, as expected for a protogyne, but were more male biased in southern regions of the GBR (Mackay and Storm Cay) than in northern regions (Lizard Island and Townsville) across all lengths and ages. The proportion of vitellogenic females also increased with length and age. Female P. leopardus were capable of daily spawning during the spawning season, but on average spawned every 4·3 days. Mature females spawned most frequently on Townsville reserve reefs (every 2·3 days) and Lizard Island fished reefs (every 3·2 days). Females on Mackay reefs open to fishing showed no evidence of spawning over 4 years of sampling, while females on reserve reefs spawned only once every 2–3 months. No effect of length on spawning frequency was detected. Spawning frequency increased with age on Lizard Island fished reefs, declined with age on Storm Cay fished reefs, and declined with age on reserve reefs in all regions. It is hypothesized that the variation in P. leopardus sex ratios and spawning frequency among GBR regions is primarily driven by water temperature, while no‐take management zones influence spawning frequency depending on the region in which the reserve is located. Male bias and lack of spawning activity on southern GBR, where densities of adult P. leopardus are highest, suggest that recruits may be supplied from central or northern GBR. Significant regional variation in reproductive traits suggests that a regional approach to management of P. leopardus is appropriate and highlights the need for considering spatial variation in reproduction where reserves are used as fishery or conservation management tools.     

Molecular and Quantitative Genetic Differentiation in Sitobion avenae Populations from Both Sides of the Qinling Mountains

Quantitative trait differences are often assumed to be correlated with molecular variation, but the relationship is not certain, and empirical evidence is still scarce. To address this issue, we sampled six populations of the cereal aphid Sitobion avenae from areas north and south of the Qinling Mountains, and characterized their molecular variation at seven microsatellite loci and quantitative variation at nine life-history traits. Our results demonstrated that southern populations had slightly longer developmental times of nymphs but much higher lifetime fecundity, compared to northern populations. Of the nine tested quantitative characters, eight differed significantly among populations within regions, as well as between northern and southern regions. Genetic differentiation in neutral markers was likely to have been caused by founder events and drift. Increased subdivision for quantitative characters was found in northern populations, but reduced in southern populations. This phenomenon was not found for molecular characters, suggesting the decoupling between molecular and quantitative variation. The pattern of relationships between F_ST and Q_ST indicated divergent selection and suggested that local adaptation play a role in the differentiation of life-history traits in tested S. avenae populations, particularly in those traits closely related to reproduction. The main role of natural selection over genetic drift was also supported by strong structural differences in G-matrices among S. avenae populations. However, cluster analyses did not result in two groups corresponding to northern and southern regions. Genetic differentiation between northern and southern populations in neutral markers was low, indicating considerable gene flow between them. The relationship between molecular and quantitative variation, as well as its implications for differentiation and evolution of S. avenae populations, was discussed.     

Relationships of latitude,altitude, and body size to litter size and mean annual production of offspring inPeromyscus

Litter size was positively correlated with latitude and altitude but not with production of offspring or with body size in Peromyscus. Increased litter size in northern populations probably reflects shortening the breeding season by climate and not a greater mortality rate at northern latitudes compared to southern latitudes. Production of offspring was negatively correlated with body size but not with latitude, altitude, or litter size. This is probably due to larger species living longer and taking longer to mature.     

The effect of supplemental food on the growth rates of neonatal,young, and adult cotton rats (Sigmodon hispidus) in northeastern Kansas,USA

In food-limited populations, the presence of extra food resources can influence the way individuals allocate energy to growth and reproduction. We experimentally increased food available to cotton rats (Sigmodon hispidus) near the northern limit of their range over a 2-year period and tested the hypothesis that seasonal growth rates would be enhanced by supplemental food during winter and spring when natural food levels are low. We also examined whether additional food resources were allocated to somatic growth or reproductive effort by pregnant and lactating females. The effect of supplemental food on growth varied with mass and season, but did not influence the growth rates of most cotton rats during spring and winter. In winter, small animals on supplemented grids had higher growth rates than small animals on control grids, but females in spring had lower growth rates under supplemented conditions. Growth rates of supplemented cotton rats were enhanced in summer. Northern cotton rat populations may use season-specific foraging strategies, maximizing energy intake during the reproductive season and minimizing time spent foraging in winter. Adult females invest extra resources in reproduction rather than in somatic growth. Pregnant females receiving supplemental food had higher growth rates than control females, and dependent pups (≤ 1 month of age) born to supplemented mothers had higher growth rates than those born to control mothers. Increased body size seems to confer an advantage during the reproductive season, but has no concomitant advantage to overwinter survival.     

Geographic Variation of Photoperiodic Diapause but Not Adult Survival or Reproduction of the Invasive Mosquito Aedes albopictus (Diptera: Culicidae) in North America

Climate differences across latitude can result in seasonal constraints and selection on life-history characters. Because Aedes albopictus (Skuse) invaded North America in the mid-1980s, it has spread across a range of ≈14° latitude and populations in the north experience complete adult mortality because of cold winter temperatures that are absent in the south. Life-table experiments were conducted to test for differences in the adult survival and reproductive schedules of Ae. albopictus females from three populations from the northern (Salem, NJ; Springfield, IL; Eureka, MO; ≈39° N) and southern (Palm Beach, Palmetto, Tampa, FL; ≈27-28° N) extremes of the species distribution in North America. There were consistent differences between northern and southern populations in incidence of photoperiodically-induced egg diapause. Under short daylength, diapause eggs constituted twice the proportion of total viable eggs from northern females (81.9-92.1%) than southern females (35.9-42.7%). There were no consistent differences between northern and southern populations in resource allocation between reproduction and maintenance, reproduction over time, and reproductive investment among offspring, and no apparent trade-offs between diapause incidence with reproduction or longevity. Our results suggest that the main response of North American Ae. albopictus to unfavorable winter climates is via the life history strategy of producing diapausing eggs, rather than quantitative variation in reproduction, and that there are no detectable costs to adult survival. Inherent geographic variation in the expression of diapause, consistent with the latitudinal extremes of A. albopictus, indicates evolutionary loss of diapause response in southern populations because of the invasion of A. albopictus in North America.     

Evidence of autumn reproduction in female European hares (Lepus europaeus) from southern Europe

Information on reproductive biology of the European hare (Lepus europaeus) in different environmental and landscape conditions comprises part of fundamental knowledge regarding species’ adaptive responses as well as many aspects of its biology. Most of the studies conducted on European hare reproduction are confined to midlatitude and northern populations, whereas no data exist on the indigenous southern populations. Here, we present information on reproductive characteristics of European hares inhabiting Mediterranean ecosystems on the island of Crete, Greece for two successive hunting seasons. Although the annual reproductive cycle of the species is well known, with an autumn sexual inactivity, the duration of this period is subjected to fluctuations in different years and for different areas. According to our data, hare populations of Crete present an autumn–early winter reproductive activity with high proportions of pregnant females observed in all the months of the study. Furthermore, the estimated mean litter size (1.54 SE ± 0.07) while signed to the lowest values ever observed for European hares is similar to values obtained in continuous breeding species of the same genus, Lepus granatensis, Lepus corsicanus, Lepus (capensis) mediterraneus, and Lepus capensis also inhabiting warm climates. In conclusion, our results suggest that Cretan European hare populations exhibit a reproductively active period during autumn–early winter where proportions of pregnant females and litter size give a strong indication of a continuous reproduction throughout the year.     

Variation in reproductive investment among and within populations of the scorpion Centruroides vittatus

Although information concerning variation among and within populations is essential to understanding an organism's life history, little is known of such variation in any species of scorpion. We show that reproductive investment by the scorpion Centruroides vittatus varied among three Texas populations during one reproductive season. Females from the Kickapoo population produced smaller offspring and larger litters than females from the Independence Creek or Decatur populations; this pattern remained when adjusting for among population variation in either female mass or total litter mass. Relative clutch mass (RCM) and within-litter variability in offspring mass (V*) did not differ among populations. Among-population variation may result from genetic differences or from phenotypically plastic responses to differing environments. Within populations, the interrelationships among reproductive variables were similar for Decatur and Independence Creek: females investing more in reproduction (measured by total litter mass, TLM) produced larger litters and larger offspring, and V* decreased with increased mean offspring mass (and with decreased litter size at Decatur). At Kickapoo, larger females produced larger litters and had larger TLM; females investing more in reproduction produced larger litters but not larger offspring. Within litter variability in offspring mass was not correlated with any reproductive variables in this latter population. These patterns may be explained by the fractional clutch hypothesis, the inability of females precisely to control investment among offspring or morphological constraints on reproduction.     

Differentiation in developmental rate across geographic regions: a photoperiod driven latitude compensating mechanism?

Genetic differentiation and phenotypic plasticity in growth rates along latitudinal gradients may benefit our understanding of latitudinal compensating mechanisms in life history patterns. Here we explore latitudinal compensatory growth mechanisms with respect to photoperiod in northern and southern populations of two damselfly species, Coenagrion puella and C. pulchellum. In addition we compared size of field‐collected adults from southern and northern populations. Eggs from females in copulating tandems were collected at two or three localities for each species in each geographic region. Eggs were transported to the laboratory and the experiment started when the eggs hatched. The role of photoperiod on the expression of larval growth rate was evaluated under controlled laboratory conditions. Both species had lower growth rate when reared in the northern photoperiod, which is counter to expectations if species use photoperiodic cues to trigger compensatory growth. Instead, both species displayed countergradient variation in growth rates, which probably enable northern populations to compensate for the shorter growth season in the north. The smaller size of field‐collected adults from northern populations also supports the view that these species compensate for the shorter growth season by investing in growth and development but accomplish this at the expense of decreased final size.     

Do seed transfer zones for ecological restoration reflect the spatial genetic variation of the common grassland species Lathyrus pratensis?

A common ecological restoration approach is the reestablishment of vegetation using seed mixtures. To preserve the natural genetic pattern of plant species local seed material should be used. Consequently, seed transfer zones (seed production areas and seed provenance regions) have been delineated for ecological restoration in Germany. Although it is assumed that these transfer zones represent genetic variation, there remains a lack of empirical data. In this study, we analyzed whether seed transfer zones reflect the genetic variation of the common grassland species Lathyrus pratensis. We sampled 706 individuals from 37 populations in Bavaria, Germany and analyzed genetic variation using amplified fragment length polymorphisms. In our study, we observed higher levels of genetic variation and fragment rarity in the southern Bavarian populations compared to northern populations. Our analyses revealed a strong genetic differentiation between southern and northern Bavarian populations delineated along the Danube River. However, seed production areas and seed provenance regions reflected genetic variation of L. pratensis only to a limited degree. Our study illustrates that the level of genetic variation within populations strongly depends on population history. Furthermore, the geomorphological and climatic attributes, which have been used to delineate seed provenance regions, do not reduce gene flow among populations. Seed collections for gene banks and seed production should comprise seeds from populations in southern and northern Bavaria representing the strong genetic variation between both regions, but prioritize southern populations due to higher levels of variation.     

Reproduction of the bank vole, Clethrionomys glareolus, in northern and southern Sweden during several seasons and in different phases of the vole population cycle

The breeding biology of the bank vole was studied in northern and southern populations in Sweden in the years 1973–76. Trapped specimens supplied organ weights and histological data, which were utilized to describe the breeding season, number and size of litters, reproductive losses, spermatogenesis, and sexual maturation. In the north the populations are cyclic. During the increase phase of the cycle the breeding season was long, litter sizes and gonad weights were large, and there was maturation of yearlings in the summer. During the peak phase reproduction was impaired in all these respects. In southern Sweden there was no cyclicity, and the reproductive events varied less, but in 1976 the litters were larger than usual and the season was much extended.
In the south the voles born during the early part of the season regularly became sexually mature during their first summer. The differences in reproductive biology are related to population dynamics and discussed in terms of different hypotheses concerning vole population cyclicity.     

Predator control and the density and reproductive success of grouse populations in Finland

The impact of predator removal or protection on the reproductive success and density of grouse populations was studied experimentally in southern and northern Finland. In predator removal areas small and medium-sized carnivores wore efficiently hunted, while in predator protection areas hunting was prohibited. Both predator (red fox, raccoon dog, pine marten and stoat) and grouse populations were monitored. The breeding success of grouse, indicated by the young/adults ratio, decreased in the predator protection area in northern Finland during a vole low. Despite a decline in the vole population, the breeding success of grouse did not decline in the predator removal area. The mean brood size during the experiment was significantly higher in the removal areas than in the protection areas both in southern and northern Finland. Predator removal/protection thus affected the reproductive success of grouse, but the impact of control on adult grouse populations was not as evident.     

Effects of latitude on the trade-off between reproduction and moult: a long-term study with pied flycatcher

Long-distance migratory passerine birds are generally time constrained by reproduction and moult, which need to be completed before migration. Breeding and post-nuptial moult may overlap especially under time-constrained conditions (northern latitudes). Here, we analysed the timing of adult moult in relation to latitude, timing of breeding and reproductive effort in pied flycatchers (Ficedula hypoleuca) breeding in four widely separated populations (40-68° N). In males but not females, the proportion of moulting birds while provisioning nestlings increased with increasing latitude. This may suggest that a moult-breeding overlap is a strategy employed by male pied flycatchers to adjust to the short breeding season at northern latitudes. However, the moult-breeding overlap was more pronounced among males in the southernmost study population (Spain). In this population, males may decide not to invest more in reproduction, and start moulting at earlier breeding stage than in northern populations,or, alternatively, birds in the Mediterranean region are time constrained by the hot and dry summer. The trade-off between breeding and post-nuptial moult may be more important in some populations than in others, depending on the latitude of the breeding site. Our results show that a moult-breeding overlap imposes a fitness cost on males in terms of fecundity and breeding success.     

Genetic Structure and Demographic History Reveal Migration of the Diamondback Moth Plutella xylostella (Lepidoptera: Plutellidae) from the Southern to Northern Regions of China

The diamondback moth Plutella xylostella (Linnaeus) (Lepidoptera: Plutellidae) is one of the most destructive insect pests of cruciferous plants worldwide. Biological, ecological and genetic studies have indicated that this moth is migratory in many regions around the world. Although outbreaks of this pest occur annually in China and cause heavy damage, little is known concerning its migration. To better understand its migration pattern, we investigated the population genetic structure and demographic history of the diamondback moth by analyzing 27 geographical populations across China using four mitochondrial genes and nine microsatellite loci. The results showed that high haplotype diversity and low nucleotide diversity occurred in the diamondback moth populations, a finding that is typical for migratory species. No genetic differentiation among all populations and no correlation between genetic and geographical distance were found. However, pairwise analysis of the mitochondrial genes has indicated that populations from the southern region were more differentiated than those from the northern region. Gene flow analysis revealed that the effective number of migrants per generation into populations of the northern region is very high, whereas that into populations of the southern region is quite low. Neutrality testing, mismatch distribution and Bayesian Skyline Plot analyses based on mitochondrial genes all revealed that deviation from Hardy-Weinberg equilibrium and sudden expansion of the effective population size were present in populations from the northern region but not in those from the southern region. In conclusion, all our analyses strongly demonstrated that the diamondback moth migrates within China from the southern to northern regions with rare effective migration in the reverse direction. Our research provides a successful example of using population genetic approaches to resolve the seasonal migration of insects.     

Let's mate here and now – seasonal constraints increase mating efficiency

1. Latitudinal climatic conditions shape the length of the mating season and could thus influence reproductive traits. Knowledge of how animals behave along latitudinal clines will increase understanding of the impact of climate on sexual selection and might help in the prediction of whether peripheral populations will spread or shrink in response to changes in climate. 2. This study investigated variation in the mating efficiency of a temperate insect, the emerald damselfly Lestes sponsa, under semi‐natural field conditions along a latitudinal gradient covering three regions of the species' distribution: south, central and north. 3. A comparison was done of the proportion of copulating males, the proportion of males that formed tandems but did not copulate (unsuccessful males), and the proportion of males that did not attempt to form a tandem (passive males) in these three regions. 4. It was found that the proportion of copulations was significantly higher at northern latitudes than in the southern and central regions. Southern latitudes had a higher proportion of successful copulations compared with central latitudes. The northern region had a significantly lower frequency of passive males. The southern region had an intermediate proportion of passive males, and the central region had the highest proportion. The proportion of unsuccessful males did not differ between regions. The population density across sites did not affect these results. 5. The study shows that damselflies inhabiting northern populations mate more intensively than individuals from southern and central populations. This suggests that more restrictive environmental conditions during a brief mating season select for higher mating efficiency.     

Trade-off between reproduction and moult – a comparison of three Fennoscandian pied flycatcher populations

Organisms that reproduce at high latitudes are assumed to have evolved several adaptations to the short summer. For birds, and especially for long-distance migrants, there is a time constraint because both reproduction and moult must be completed before autumn migration. It has therefore been assumed that birds at northern latitudes must initiate their moult during reproduction more often than birds at low latitudes. To investigate how passerine birds breeding at different latitudes allocate their time between reproduction and moult, we compared timing of these activities during three consecutive breeding seasons in three widely separated populations of the pied flycatcher Ficedula hypoleuca. Our results show that the frequency of individuals with moult-breeding overlap, and moult initiation in relation to breeding stage, varied considerably among populations and years. In all three populations, female moult initiation was restricted to the late nestling period. The males had a more pronounced moult-breeding overlap than the females, but its duration was similar in all three study areas. Thus, there was no evidence for a more pronounced moult-breeding overlap at high compared with low latitudes. These results suggest that pied flycatchers sometimes accept a moult-breeding overlap, but that the time gained by having too extensive an overlap between reproduction and moult does not outweigh the associated costs. Long-distance migrants breeding at northern latitudes apparently experience a trade-off between reproduction and somatic investment during moult. We therefore suggest that a pronounced moult-breeding overlap is not a typical strategy used by long-distance migrants to adjust to the short breeding season at northern latitudes. Received: 7 May 1998 / Accepted: 24 August 1998     

Influence of seasonal time constraints on growth and development of common frog tadpoles: a photoperiod experiment

Organisms living in seasonal environments are often limited by the time available to complete their development. Especially individuals in northern populations may face severe time constraints in their need of completing development before the end of the growth season. Larval amphibians have been widely used in studies of phenotypic plasticity. However, their responses to changes in photoperiod, the main seasonal cue in many organisms, are unknown. In a laboratory experiment, we studied whether common frog (Rana temporaria) tadpoles originating from two populations (separated latitudinally by 1600 km) adjust their growth and development according to the progress of the season by using photoperiodic cues, and whether these responses are temperature dependent. We hypothesised that if frogs use photoperiod as a cue, they should increase growth and development rates as a response to photoperiodic treatments mimicking progressing season. Although our predictions were not verified in either of the populations, photoperiod manipulations had effects on larval life history in both populations. When exposed to progressing season treatments and high temperature, tadpoles from the southern population ceased feeding, which led to delayed metamorphosis and increased mortality. In the northern population, age at metamorphosis was unaffected by the photoperiod treatments, but growth rate until metamorphosis and metamorphic size were slightly larger in the treatments with shorter (increasing or decreasing) day length. Irrespective of photoperiod treatments, growth and development rates, size at metamorphosis and food consumption were higher in the northern as compared to the southern population. These results indicate that in contrast to several insect species, the critical life history decisions in amphibian larvae may not be strongly influenced by photoperiodic cues, but different populations seem to differ in this respect. However, the strong temperature×photoperiod interactions in several traits in the southern population suggest that the role of photoperiodic cues may be affected by other environmental factors, although the ecological significance of these differences remains unclear.     

Effects of photoperiodically induced reproductive diapause and cold hardening on the cold tolerance of Drosophila montana

Coping with seasonal and daily variation in environmental conditions requires that organisms are able to adjust their reproduction and stress tolerance according to environmental conditions. Females of Drosophila montana populations have adapted to survive over the dark and cold winters at high latitudes and altitudes by spending this season in photoperiodically controlled reproductive diapause and reproducing only in spring/summer. The present study showed that flies of a northern population of this species are quite tolerant of low temperatures and show high seasonal and short-term plasticity in this trait. Culturing the flies in short day length (nearly all females in reproductive diapause), as well as allowing the flies to get cold hardened before the cold treatment, increased the cold tolerance of both sexes both in chill coma recovery time test and in mortality assay. Chill coma recovery time test performed for the females of two additional D. montana populations cultured in a day length where about half of the females enter diapause, also showed that diapause can increase female cold tolerance even without a change in day length. Direct linkage between diapause and cold tolerance was found in only two strains representing a high-altitude population of the species, but the phenomenon will certainly be worth of studying in northern and southern populations of the species with larger data sets.     

Effect of Temperature on Rearing of the First Brood by the Founder Females of the Ant Lasius niger (Hymenoptera, Formicidae) : Latitude-Dependent Variability of the Response Norm

It has been established that duration of brood development and success of rearing of the first worker individuals by founder females of the ant Lasius niger from the northern population (St. Petersburg) depend on temperature conditions to a larger extent than of individuals from the southern areal (Borisovka Village, Belgorod Region) The development rate of individuals from the northern population is lower as compared with individuals from the southern population in the low temperature range (17–20°C), but is higher at elevated temperatures (22–27°C). This leads to a greater slope of the temperature regression line of the development rate on the temperature, to lower values of the sum of effective temperatures, and to a higher temperature threshold of development in the northern population. Lower temperatures have a stronger effect on development of larvae from the northern population by inducing diapause in the greater number of individuals as compared with larvae from the southern population. The L. niger founder females require, on average, higher temperatures than the females from the southern populations of this species for rearing the first worker ants during the same summer season (i.e., without winter diapause). Thus, ants of the southern populations, when rearing the first brood, use a strategy of faster development at higher temperatures, while low temperatures restrict to the greater extent the development of their brood as compared with ants from the southern populations. It is concluded that one of the ways of physiological adaptation of ants for inhabitance on the North is such change of the norm of response to temperature, at which development becomes more temperature-dependent, the physiological response to higher temperatures increasing at the cost of its decrease at lower temperatures.