Diurnal changes in phosphoenolpyruvate carboxylase and pyruvate, orthophosphate dikinase properties in the natural environment: interplay of light and temperature in a C4 thermophile

Activities of phosphoenolpyruvate (PEP) carboxylase (EC 4.1.1.31) were measured in leaf extracts of field grown Amaranthus paniculatus L. (C₄) during a natural diurnal irradiance and temperature pattern. Enzyme assays were run at both fixed (30°C) and the corresponding leaf temperature at the time of harvest. Light activation of PEP carboxylase (PEPCase) at fixed assay temperatures was expressed as a decrease in S_0–5 (PEP) after a threshold (> 330 μmol m^–2 s^–1) photon fluence rate was surpassed at noon. Earlier in the morning, increase in apparent enzyme affinity for PEP was observed when the assay was run at leaf temperature, indicating a physiologically meaningfull effect of temperature on S^0.5 (PEP). The 3.3-fold increase in PEPCase activity at low PEP and fixed assay temperature between the minimal and maximal irradiance and temperature hours of the day, became 12.8-, 11.5- and 7.4-fold when assays were run at the corresponding leaf temperature during three diurnal cycles with respective temperature differences (max minus min) of 9.0, 8.3 and 7.4°C. The extent of malate inhibition was the same for both day and night forms of PEPCase assayed at 35°C, but increased considerably with night enzyme at 25°C. The results indicate that light increases the apparent affinity of PEPCase for PEP and that at lower temperatures malate becomes more inhibitory. Pyruvate orthophosphate dikinase activity started to increase immediately after sunrise and the 10-fold increase at fixed temperature became 14.8-, 14.2- and 13.1-fold when assays were run at the above leaf temperatures. This indicates that the light effect predominates with pyruvate, orthophosphate dikinase, while with phosphoenolpyravate carboxylase, light and temperature co-operate to increase the day enzyme activities.     

Thermal ecology and spatio-temporal distribution of the Mediterranean lizard Psammodromus algirus

We studied summer activity patterns and thermoregulation in the Mediterranean lizard Psammodromus algirus in a holm oak wood in Central Spain. The circadian rhythm curve was bimodal, with a pronounced peak after sunrise, a minimum at noon and a second lower peak in the afternoon. Increased activity in the morning could be explained by insolation levels, whereas the midday low was caused by the rise in temperature.
There was a strong positive correlation between body temperature and both air and ground temperatures. Nevertheless, there was no significant correlation between body and air temperatures when the effect of radiation was removed, which was done by considering only individuals in the shade at high temperatures.
Activity showed a bell-shaped relationship with body and air temperatures, with maximum activity at 24.5°C (air temperature) and 31.4°C (body temperature). The spatial distribution pattern of P. algirus (selection of sunny areas and distance to the vegetation edge) reflects the environmental temperatures that are optimal for activity; the lizards show a strong preference for sunny patches when air temperature is lower than the optimum. We conclude that heliothermy is the only viable thermoregulation pattern in Mediterranean forest environments, with heat conduction playing a negligible rote.     

High and low temperature tolerances and their relationships to distribution of agaves

Abstract. The potential influence of tissue tolerances to extreme temperatures on distributional limits was investigated for 15 taxa (14 species) of leaf-succulent agaves from the south-western United States and northern Mexico. As a group, the agaves exhibited a moderate low temperature tolerance of – 11°C (based on a 50% inhibition in the number of mesophyll cells taking up a stain, neutral red). However, nearly all of the species were able to tolerate extremely high tissue temperatures of over 60°C. Nocturnal acid accumulation by these crassulacean acid metabolism plants was about 6°C more sensitive to temperature extremes than was cellular membrane integrity.
High and low temperature acclimation in response to changing day/night air temperatures was observed in all 15 taxa, with high temperature acclimation averaging two-fold greater than low temperature acclimation (3.8°C versus 2.0°C per 10°C change in ambient temperature). Species occupying the coldest habitats exhibited the greatest low temperature tolerances and acclimation; several such species, such as Agave utahensis and A. schottii , had small rosette sizes which resulted in higher minimum leaf temperatures. Species from the hottest habitats had among the greatest high temperature tolerances and acclimation; the two species from open desert scrub habitats, A. deserti and A. lecheguilla , had the lowest leaf shortwave absorptances observed, which would result in lower maximum leaf temperatures. Thus morphology and tissue tolerances to stressful temperatures reflect the temperature extremes of a plant's native habitat, although low temperature tolerance appears to limit the distribution of agaves more than high temperature tolerance.     

Shoot temperatures and thermal tolerances for succulent species of Haworthia and Lithops

Abstract. Temperatures of small succulent plants, such as species in the genera Haworthia and Lithops , are highly influenced by temperatures of the surrounding soil. Indeed, the minimum and the maximum temperatures of the upper leaf epidermis of Haworthia retusa. H. turgida. Lithops leslei , and L. turbiniformis were generally within 1°C of the accompanying soil surface temperatures. An energybudget model closely predicted such soil-to-plant temperature differences as well as the effect of the greater convective exchange for the protruding Haworthia species compared with the Lithops species, which were flush with the soil surface. Although a lower shortwave absorptance would reduce maximum shoot temperatures, the shortwave absorptances of all four species were similar to those of the soil in their respective native habitats in South Africa. Tolerances of the four species to low and to high temperatures at three different day/night air temperatures (15°C/5°C, 30°C/20°C, and 45°C/35°C) were analysed using cellular accumulation of a vital stain, neutral red. Chlorenchyma cells were slightly more tolerant of extreme temperatures than were cells of the water-storage parenchyma. In this regard, H. retusa survived low and high temperatures that killed the water-storage parenchyma but not the chlorenchyma. Acclimation to low temperatures and to high temperatures, which was exhibited by all four species, led to estimated tolerances to 1 h at −16°C and 68°C. Although the low temperature tolerance is not particularly noteworthy, very few vascular plants are reportedly able to tolerate such high temperatures.     

A field study of photosynthetic temperature acclimation in Carex eleocharis Bailey

Abstract. Seasonal patterns in photosynthetic temperature acclimation and growth were investigated in the sedge, Carex eleocharis Bailey, a species which has demonstrated a marked capacity for shifts in the photosynthetic temperature optimum in previous growth chamber studies. The seasonal production of new leaves was 90% complete by the earliest study date, June 3. Shifts in the photosynthetic temperature optimum of 10°C (from 15 to 25°C) were observed during the months of June and July. These results indicate that in situ acclimatory adjustments in C. eleocharis occur in existing leaf tissue, rather than new leaves which are produced as the season progresses. Despite the 10°C increase in the temperature optimum, mean mid-day leaf temperatures were higher than the optimum throughout the summer. A broad temperature response appeared to be more important than the acclimation adjustments in maintaining near-maximum photosynthesis rates during the mid-day period. Seasonal shifts in the photosynthetic temperature optimum were not as great as those previously observed in growth chamber studies. This discrepancy arises because of the capacity for growth chamber grown plants to produce new leaves with temperature response characteristics closely tuned to the growth temperature regime. In field-grown plants the production of 90% of the leaves during the cool portion of the season places limitations on the potential for acclimation to the warmer midsummer temperatures.     

The effect of flutter on the temperature of poplar leaves and its implications for carbon gain

The leaf temperatures of two poplar species (Populus tremuloides Michx. and P. fremontii Wats.) were characterized by attaching thermocouples to leaves that were either constrained to a fixed position or allowed to flutter naturally. There were no observed temperature differences between fluttering and constrained leaves in the lower canopy, but fluttering leaves at the top of the canopy were as much as 2–4°C cooler than constrained leaves. An increase in heat transfer, a decrease in light interception or both could account for these observed differences in the temperature of fluttering versus constrained leaves. Fluttering can increase the boundary-layer conductance to convective heat exchange by as much as 50 and 20% for laminar and turbulent flow, respectively. The benefit that these leaf temperature differences may provide to the carbon economy of a poplar canopy was dependent on the ambient temperature. Populus fremontii, which is frequently exposed to daytime temperatures exceeding 35°C during summer months in the central valley of California, USA, could show an increase in carbon gain as a result of lower upper canopy leaf temperatures. For aspen, the benefit would be much smaller and often negative because of much lower air temperatures. Lower leaf temperatures may also increase the water use efficiency of poplars. However, the maintenance of lower leaf temperatures may not be the primary adaptive significance of leaf flutter.     

Effects of temperature on the growth of spring ephemerals: Crocus vernus

Spring ephemerals of deciduous forests appear shortly after snow melt and senesce shortly after the overstorey canopy has closed. During this short period, they take advantage of the high light conditions, but also of the low-temperature regime that favours growth in these species. This better growth at cooler temperatures has been attributed to longer leaf life duration. The objective of this study was to test if temperature effect on growth in spring ephemerals can be explained solely by the leaf life duration, or if there is also a direct temperature effect on the underground organ growth. Crocus vernus (L.) Hill was exposed to two temperature regimes, 12/8 and 18/14°C (day/night), for sequential harvests and to four differential air to soil temperature regimes: 12/12, 12/18, 18/12 and 18/18°C (air/soil temperature). Dry mass, leaf nitrogen, starch concentration and cell size of the new corm were determined. Final dry mass and cell size were higher at the lower temperature regime while leaf lasted longer than at the higher temperature regime. Although both air and soil temperatures had an impact on the different growth parameters measured, the impact of soil temperatures was greater and it also influenced leaf life duration. At higher temperature, corm growth stopped before the first visual sign of leaf senescence, suggesting that corm growth controls leaf life duration in C. vernus rather than the opposite and that crocus growth becomes rapidly sink limited at higher temperatures.     

Chilling sensitivity of the frost-tolerant potato Solanum commersonii

Frost tolerance has been reported in the shoots of wild, tuberiferous potato species such as Solanum commersonii when the plants are grown in either field or controlled conditions. However, these plants can survive as underground tubers and avoid unfavorable environmental conditions altogether. As such, leaf growth and photosynthesis at low temperature may not be required for survival of the plants. In order to determine the temperature sensitivity of S. commersonii shoots, we examined leaf growth, development and photosynthesis in plants raised at 20/16°C (day/night). 12/9°C and 5/2°C. S. commersonii leaves grown at 5°C exhibited a marked decrease in leaf area and in total chlorophyll (Chl) content per leaf area when compared with leaves grown at 20°C. Furthermore, leaves grown at 5°C did not exhibit the expected decrease in either water content or susceptibility to low-temperature-induced photoinhibition that normally characterizes cold acclimation in frost-tolerant plants. Measurements of CO₂-saturated O₂ evolution showed that the photosynthetic apparatus of 5°C plants was functional, even though the efficiency of photosystem II photochemistry was reduced by growth at 5°C. A decrease in the resolution of the M-peak in the slow transients for Chl a fluorescence in leaves grown at 12 and 5°C and in all leaves exposed to high light at 5°C indicated that low temperature significantly affected processes on the reducing side of Q_A, the primary quinone electron acceptor in photosystem II. Thus S. commarsonii exhibits the characteristics of a plant that is limited by chilling temperatures. Although S. commersonii can tolerate light frosts, its sensitivity to chilling temperatures may result in shoot dieback in winter in its native habitat. The plants may avoid both chilling and freezing temperatures by overwintering as underground tubers.     

Thermal ecology of the mudskippers, Periophthalmus koelreuteri (Pallas) and Boleophthalmus boddarti (Pallas) of Kuwait Bay

The annual range of body temperatures (14–35°C) of emergent mudskippers are substantially less than that of air temperatures (10–42°C) as a result of behavioural thermoregulation. In winter, low surface temperatures are avoided by remaining in burrows. Newly emerged mudskippers then bask until body temperatures rise above 14°C before they move onto the mud. In summer, body temperatures are kept lower than ambient by selecting areas where evaporative cooling is high. Body temperatures generally match those of wet mud, which can be 7°C lower than air shade temperatures. The smaller, more terrestrial, Periophthalmus koelreuteri have body temperatures which are mainly lower in summer and higher in winter than Boleophthalmus boddarti .     

Short-Term Changes in Heat Tolerance in the Alpine Cushion Plant Silene acaulis ssp. excapa [All.] J. Braun at Different Altitudes

Abstract: The habit of cushion growth positively affects plant temperature but at the same may increase the risk of occasional overheating. In order to determine the adaptive response to short-term heat stress, we exposed S. acaulis cushions at field sites to controlled heat treatments using infrared lamps. Natural diurnal changes in heat tolerance were monitored at alpine sites and at a site distinctly below the natural distribution boundary, where higher temperatures were expected. The range of heat tolerance limits in summer, 45.5 - 54.5 °C (9 K), exceeded that reported for other alpine species (0.1 - 5 K) and even that for total seasonal changes (5 - 8 K). Heat tolerance either increased or decreased on most days (80 %). The maximum diurnal increase was + 4.7 K. Under the experimental conditions heat hardening started at leaf temperatures around 30 °C and proceeded at mean rates of 1.0 ± 0.5 K/h. The onset of functional disturbances in photosystem II also occurred at 30 °C. Heating rates exceeding those naturally found above 30 °C (> 10 K/h) appeared to retard heat hardening. During summer average leaf temperature maxima were 12.4 K (600 m) and 13.0 K (1945 m) higher than air temperature which corroborates the heat trapping nature of cushion plants. At 600 m, as compared to 1945 m, cushions experienced significantly higher leaf temperature maxima (+ 8.8 K) and exceeded 30 °C on most days (80 %). This resulted in a significantly higher heat tolerance (LT₅₀) at 600 m (51.7 ± 0.2 °C) than at 1945 m (49.8 ± 0.2 °C). The fast short-term changes of heat tolerance in summer help S. acaulis to cope with the occasional diurnal short-term heat stress associated with cushion growth.     

MICROCLIMATIC EFFECTS ON WATER RELATIONS,LEAF TEMPERATURES,AND THE DISTRIBUTION OF HERACLEUM LANATUM AT HIGH ELEVATIONS

The small-scale distribution of an understory herb, Heracleum lanatum, was evaluated in terms of leaf temperature and water relations limitations due to a large leaf size (630 cm²). Diurnal variations in transpiration (4 to 60 mg m⁻² s⁻¹) were influenced by fluctuations in solar irradiance, wind speed, leaf temperature and stomatal conductance. Computer simulations indicated that leaf temperatures in a forest clearing would be > 12 C above air temperature, with maximum transpiration rates of 140 mg m⁻² s⁻¹, and daily water loss to be over 200% greater than values at natural understory locations. Simulations of nocturnal temperature relations indicated ~100 W m ⁻² less incident longwave irradiance in the forest clearing as compared to the understory (560 vs. 660 W m⁻² at 400 hr). This difference led to predicted leaf temperatures being as low as 6 C below air temperature in the forest clearing while measured leaf temperatures in the forest understory were within 1.5 C of air temperature throughout the night. Furthermore, minimum air temperatures were at or below 6 C on 36% of the nights during the summer growth period indicating that in open areas leaves of H. lanatum would frequently be below 0 C and subject to possible freeze damage. Heracleum lanatum may be more abundant in the shaded understory of the subalpine forest because exposure in open environments would result in high leaf temperatures and increased transpirational water loss during the day, as well as low leaf temperatures with the possibility of freeze damage at night.     

Analysis of the response of leaf extension to chilling temperatures in Lolium temulentum seedlings

Lolium temulentum L. plants were grown at 20°C and transferred to 2°C or 5°C at 21, 28 or 35 days after sowing, when leaves 3, 4 and 5, respectively, were at mid-expansion and leaves 4, 5 and 6 were just emerging. Leaves of plants exposed to 2°C for 7 or 14 days before their date of emergence at 20°C failed to appear at all during the course of the experiment. Transfer to 2°C at emergence resulted in a delay of about 40 days before expansion was detected and subsequent growth was extremely slow. By contrast, although leaves of seedlings exposed to 5°C at or prior to emergence were significantly smaller and slower-growing than the same leaves of plants maintained at 20°C, the difference in vegetative development and tillering between 2°C and 5°C was much less marked than between 5°C and 2°C, implying the existence of a rather sharp threshold for growth between the latter temperatures. Leaves transferred to 2°C at mid-expansion attained a final size not very different from leaves exposed to 5°C at the same time, but expension rates were only 20–30% of those at 5°C, and the time taken to achieve full expansion a corresponding 3 to 5 times longer. These responses were quantified by fitting Richards functions to measurements of leaf extension and determining, from the parameters of the curves, asymptotic maximal lengths, mean relative and absolute extension rates, inflexion points and durations of growth. The potential usefulness of Lolium temulentum as a model species for studying the relationship between temperature and growth in the Granmineae is discussed.     

Temperature and water relation patterns in subalpine understory plants

Summary The daily temperature and water relations of 7 perennial subalpine, understory species (1 shrub, 1 subshrub, 5 herbs) were compared in the Rocky Mountains of southeastern Wyoming with an emphasis on the effects of natural sun and shade exposure. Field measurements of rainfall; leaf, air, and soil temperatures; stomatal conductance to water vapor diffusion; and plant and soil water potentials were supplemented with leaf and root morphological measurements to evaluate potential adaptive patterns in understory species.Morphologically, all 7 species had relatively broad leaves that were hypostomous and bicolored with the abaxial leaf surface lighter than the abaxial surface. Root systems tended to be shallow (<20 cm), especially for the herbaceous species. Although soil water potentials from 4 to 40 cm depths remained relatively high throughout the summer (>-1.0 MPa), plant xylem water potentials for sunlit plants decreased to below-2.0 MPa during midday. During these sunlit periods, leaf temperatures and conductances increased substantially, leading to severe wilting for 4 of the 5 herbaceous species. Stomatal conductance and density for 6 of the 7 species monitored were much greater on abaxial compared to adaxial leaf sides and substantial stomatal closure occurred when either leaf side was oriented to receive direct sunlight. Moreover, stomatal opening on abaxial leaf sides corresponded to the amount of sunlight incident upon the adaxial rather than abaxial leaf surfaces. The 2 shrubby species did not wilt during these periods and were characterized by the highest leaf temperatures (>30°C). These 2 species also had consistantly lower xylem water potentials throughout the summer growth period. These results are discussed in terms of the possible adaptive significance of midday wilting, leaf hypostomy and bicoloration and stomatal behavior to the water and photosynthetic relations of understory species.     

The effects of temperature on germination and early growth of three plant species indigenous to Central Australia

The temperature responses of the perennial grass Eragrostis eriopoda Benth., an all-season annual grass (Enneapogon polyphyllus Domin.) and a cool-season annual forb (Lepidium muelleri-ferdinandi Thell.) were studied in two experiments, one during germination and the other during the establishment phase. In the first experiment, extreme differences in the response of germination to temperature occurred between species. The optimum was 42°C for Eragrostis and 15 to 21° C for Enneapogon and Lepidium although Enneapogon tolerated higher temperatures than did Lepidium. The effect of temperature on speed of germination is discussed. In the second experiment, seedlings were assigned to six day/night temperature regimes ranging from 24/16 C to 45/31 C for three weeks in a phytotron. Both grasses responded positively to temperatures tip to 30/22°C with little further change up to 45/37° C, while Lepidium died at temperatures greater than 30/22° C. The experimentally determined response to temperature reflected field observations of seasonal establishment patterns for each species. The ecological significance of the results is discussed.     

Effects of water stress and nocturnal temperature on carbon allocation in the perennial grass, Leymus chinensis

Water deficit and high temperature often occur simultaneously, but their effects on plants are usually investigated separately. The aim of this study was to test how interactions between water stress and nocturnal warming affect carbon allocation in the perennial grass, Leymus chinensis . Plant biomass, dry mass allocation, ¹⁴C partitioning and carbon isotope composition (δ¹³C) were measured. Severe and extreme water stress during nocturnal warming decreased the allocation of dry mass and ¹⁴C partitioning below ground to the roots, but moderate water stress significantly increased the below-ground allocation of dry mass and ¹⁴C, especially at the lower night temperature. The δ¹³C values were more positive at day/night temperatures of 30/20°C than at 30/25°C, and greater in the roots than in the leaves. By plotting the δ¹³C values of the leaves against the δ¹³C values of the roots, the slopes of regressions were steeper at low than at high night temperature, also indicating that nocturnal warming reduces carbon allocation below ground to the roots. The results suggest that nocturnal warming may weaken acclimation during water stress in this species by regulating carbon allocation between source and sink organs.     

Shifts in the optimal temperature for nocturnal CO2 uptake caused by changes in growth temperature for cacti and agaves

When the day/night air temperatures were raised from 10°C/10°C to 30°C/30°C, the optimal tempearture for nocturnal CO² uptake by six species of cacti and three species of agave shifted from an average of 12°C to an average of 20°C. The maximum rate of CO² uptake was higher for Agave americana at the higher ambient temperature, lower for A. deserti , and much lower for A. utahensis , consistent with the relative mean temperatures of their native habitats. For the cactus Coryphantha vivipara , which had the greatest temperature shift observed (13°C), the halftime was 8 days for the upward shift and 4 days for the downward shift. The halftimes for the comparable shifts averaged 1.6 days for three other species of cacti and less than 1 day for two agave species. The shifts in the optimal temperature for nocturnal CO² uptake were in response to changes in nighttime temperature, at least for C. vivipara , and reflected temperature responses of both the stomates and the chlorenchyma.     

Mode of high temperature injury to wheat during grain development

High temperature stress adversely affects wheat growth in many important production regions, but the mode of injury is unclear. Wheat ( Triticum aestivum L. cv. Newton) was grown under controlled conditions to determine the relative magnitude and sequences of responses of source and sink processes to high temperature stress during grain development. Regimes of 25°C day/15°C night, 30°C day/20°C night, and 35°C day/25°C night from 5 days after anthesis to maturity differentially affected source and sink processes. High temperatures accelerated the normal decline in viable leaf blade area and photosynthetic activities per unit leaf area. Electron transport, as measured by Hill reaction activity, declined earlier and faster than other photosynthetic processes at the optimum temperature of 25/15 °C and at elevated temperatures. Changes in RUBP carboxylase activities were similar in direction but smaller in magnitude than changes in photosynthesic rate. Increased protease activity during senscence was markedly accentuated by high temperature stress. Specific protease activity increased 4-fold at 25/15 °C and 28-fold at 35/25 °C from 0 to 21 days after initiation of temperature treatments. Grain-filling rate decreased from the lowest to the highest temperature, but the change was smaller than the decrease in grain-filling duration at the same temperatures. We concluded that a major effect of high temperature is acceleration of senescence, including cessation of vegetative and reproductive growth, deterioration of photosynthetic activities, and degradation of proteinaceous constituents.     

Thermoregulatory behaviour, diel activity and the relationship of spontaneous locomotor activity to temperature in the sea raven, Hemitripterus americanus (Gmelin)

Twelve sea ravens, Hemitripterus americanus , were tested individually for three day periods in electronic shuttleboxes to measure their preferred and avoided temperatures, diel activity pattern and relationship of activity to temperature. The fish avoided temperatures below 9°C or above 25°C, preferring temperatures with a central tendency of 15–17°C. The fish were primarily nocturnal, being more active at night than by day. Locomotor activity increased eightfold between 10 and 20°C, but exhibited an anomalous decrease between 14 and 18°C, corresponding to the preferred-temperature zone, characteristic of maximal summer temperatures in the natural habitat.     

LEAF SURFACE WETNESS AND GAS EXCHANGE IN THE POND LILY NUPHAR POLYSEPALUM (NYMPHAEACEAE)

As part of a continuing study of the effects of leaf surface wetness on gas exchange, the occurrence of leaf surface wetting by dewfall and associated effects on photosynthesis were evaluated for floating and aerial leaves of the pond lily Nuphar polysepalum Engelm. Because of nighttime radiation exchange with a cold sky, high humidity, and the presence of adaxial stomata, we predicted that pond lily leaves would be particularly susceptible to wetting events such as dewfall. A substantial reduction in net photosynthesis (up to 20%) occurred for leaves that were experimentally misted to simulate leaf wetting by dewfall. Aerial leaves remained below dewpoint temperatures for long periods on clear nights. However, floating leaves rarely approached dewpoint temperatures at night because minimum nighttime temperatures of leaves were up to 10 C warmer than air temperature. Thus, floating leaves of N. polysepalum did not experience dew formation primarily because of strong thermal coupling to a substrate (water) that was much warmer than air temperature at night. This coupling to a warmer substrate prevented a potentially strong inhibition of photosynthetic CO₂ exchange the following morning.