Effects of elevated atmospheric CO(2) and temperature on leaf optical properties in Acer saccharum

Elevated partial pressures of atmospheric carbon dioxide, similar to numerous causes of plant stress, may alter leaf pigmentation and structure and thus would be expected to alter leaf optical properties. Hypotheses that elevated CO(2) pressure and air temperature would alter leaf optical properties were tested for sugar maple (Acer saccharum) in the middle of its fourth growing season under treatment. The saplings had been growing since 1994 in open-top chambers and partial shade at Oak Ridge, Tennessee under the following treatments: (1) ambient CO(2) pressure and air temperature (control); (2) CO(2) pressure approximately 30 Pa above ambient; (3) air temperatures 3 degrees C above ambient; and (4) elevated CO(2) and air temperature. Under elevated CO(2) or temperature, spectral reflectance, transmittance and absorptance in the visible spectrum (400-720 nm) tended to change in patterns that generally are associated with chlorosis, with maximum differences from the control near 700 nm. However, these changes were not significant at P=0.05. Although reflectance, transmittance and absorptance at 700 nm correlated strongly with leaf chlorophyll concentration, variability in chlorophyll concentration was greater within than among treatments. The lack of treatment effects on pigmentation explained the non-significant change in optical properties in the visible spectrum. Optical properties in the near-infrared (721-850 nm) were similarly unresponsive to treatment with the exception of an increased absorptance throughout the 739-850 nm range in leaves that developed under elevated air temperature alone. This response might have resulted from effects of air temperature on leaf internal structure.     

Multi-objective environment chamber system for studying plant responses to climate change

This paper describes the technical information and performance of a new multi-objective chamber system enabling the control of environmental variables (e.g., temperature, CO₂, air humidity, wind speed, and UV-B radiation) for understanding plant responses to climate change. Over a whole growing season, four different climate scenarios were evenly programmed into the system’s 16 chambers as ambient environment (AMB), elevated temperature (ET), elevated CO₂ concentration (EC) and elevated temperature and CO₂ concentration (ETC). Simultaneously, the chamber effects were assessed regarding the physiological responses and growth of a boreal perennial grass (reed canary grass, Phalaris arundinacea L.). During the growing season, the chamber system provided a wide variety of climatic conditions for air temperature (T _a), relative humidity (RH) and CO₂ concentration (C _a) in the AMB chambers following outside conditions. The target temperature (+3.5°C) was achieved to a good degree in the ET and ETC chambers, being on average 3.3°C and 3.7°C higher than ambient conditions, respectively. The target concentration of CO₂ (700 ppm) was also well achieved in the EC and ETC chambers, being on average 704 ppm and 703 ppm, respectively. The stable airflow condition inside all of the chambers provided a homogeneous distribution of gases and temperature. The decreases in RH and increases in vapour pressure deficit (VPD) in the elevated temperature chambers were also maintained at a low level. Chamber effects were observed, with some physiological and growth parameters of plants being significantly lower in the AMB chambers, compared to outside conditions. The plant growth was negatively affected by the reduced radiation inside the chambers.     

Effects of climate warming and species richness on photochemistry of grasslands

In view of the projected climatic changes and the global decrease in plant species diversity, it is critical to understand the effects of elevated air temperature (T_air) and species richness (S) on physiological processes in plant communities. Therefore, an experiment of artificially assembled grassland ecosystems, with different S (one, three or nine species), growing in sunlit climate-controlled chambers at ambient T_air and ambient T_air + 3°C was established. We investigated whether grassland species would be more affected by midday high-temperature stress during summer in a warmer climate scenario. The effect of elevated T_air was expected to differ with S. This was tested in the second and third experimental years by means of chlorophyll a fluorescence. Because acclimation to elevated T_air would affect the plant's stress response, the hypothesis of photosynthetic acclimation to elevated T_air was tested in the third year by gas exchange measurements in the monocultures. Plants in the elevated T_air chambers suffered more from midday stress on warm summer days than those in ambient chambers. In absence of severe drought, the quantum yield of PSII was not affected by elevated T_air. Our results further indicate that species had not photosynthetically acclimated to a temperature increase of 3°C after 3 years exposure to a warmer climate. Although effects of S and T_air × S interactions were mostly not significant in our study, we expect that combined effects of T_air and S would be important in conditions of severe drought events.     

A whole-tree chamber system for examining tree-level physiological responses of field-grown trees to environmental variation and climate change

A whole-tree chamber (WTC) system was installed at Flakaliden in northern Sweden to examine the long-term physiological responses of field-grown 40-year-old Norway spruce trees [Picea abies (L.) Karst.] to climate change. The WTCs were designed as large cuvettes to allow the net tree-level CO(2) and water fluxes to be measured on a continuous basis. A total of 12 WTCs were used to impose combinations of atmospheric carbon dioxide concentration, [CO(2)], and air temperature treatments. The air inside the ambient and elevated [CO(2)] WTCs was maintained at 365 and 700 micromol mol(-1), respectively. The air temperature inside the ambient temperature WTCs tracked air temperature outside the WTCs. Elevated temperatures were altered on a monthly time-step and ranged between +2.8 and +5.6 degrees C above ambient temperature. The system allowed continuous, long-term measurement of whole-tree photosynthesis, night-time respiration and transpiration. The performance of the WTCs was assessed using winter and spring data sets. The ability of the WTC system to measure tree-level physiological responses is demonstrated. All WTCs displayed a high level of control over tracking of air temperatures. The set target of 365 micromol mol(-1) in the ambient [CO(2)] chambers was too low to be maintained during winter because of tree dormancy and the high natural increase in [CO(2)] over winter at high latitudes such as the Flakaliden site. Accurate control over [CO(2)] in the ambient [CO(2)] chambers was restored during the spring and the system maintained the elevated [CO(2)] target of 700 micromol mol(-1) for both measurement periods. Air water vapour deficit (VPD) was accurately tracked in ambient temperature WTCs. However, as water vapour pressure in all 12 WTCs was maintained at the level of non-chambered (reference) air, VPD of elevated temperature WTCs was increased.     

Mild experimental climate warming induces metabolic impairment and massive mortalities in southern African quartz field succulents

Current thermal regimes for many southern African succulent species of the subfamily Ruschioideae, which rapidly diversified during the cooler Pleistocene period, may be close to their tolerable extremes, and likely exceeded with anticipated future climate warming. This hypothesis was tested by exposing succulent species of different size and architecture to differently elevated temperatures approximating future African climate scenarios (2.5–3.8 °C increases in mean annual daily temperature maxima) using transparent hexagonal open-top chambers of different heights. Air temperatures, soil water potentials and amounts of fog and dew precipitation were monitored hourly in the differently heated open-top chambers and ambient environment, and changes in species leaf densities and canopy covers precisely determined in these chambers and ambient environment from high resolution digital images taken at 3-monthly intervals spanning a 12-month monitoring period. Photochemical efficiencies and activities of the photosynthetic enzyme Rubisco were also measured in one widespread dwarf succulent species following 2-h exposures of its populations in a forced draft oven to eight different heat intensities (range: 40–54 °C), the highest closely matching the temperature extreme of 54.8 °C recorded in the most intensively heated open-top chambers. After 12-months warming, all succulent species displayed massively (up to 90.2%) reduced leaf densities and canopy covers in the differently heated open-top chambers, with small sparsely branched species comprising single leaf pairs per axis exhibiting much greater reductions than large, shrubby or creeping species with multiple leaves. Noteworthy, was that fog and dew precipitation levels and soil water potentials at the centres of the least intensively heated chambers did not differ significantly from those in the ambient environment, even during the critical dry summer and early autumn seasons. However, leaf density and canopy reductions in these chambers were of similar magnitude to those in the most intensively heated chambers where fog and dew precipitation levels and soil water potentials were significantly reduced. These findings identified elevated temperatures as the principal cause of the observed massive reductions in leaf density and canopy cover and supported the hypothesis that mild anthropogenic warming could exceed the thermal thresholds of many southern African quartz field succulents leading to metabolic impairment. This impairment explained by an observed loss in the catalytic efficiency of Rubisco at daytime temperature extremes exceeding 54 °C, preceded by a decrease in PSII electron transport commencing at temperatures much lower than the threshold for Rubisco de-activation.     

Temperature magnified postcapture mortality in adult sablefish after simulated trawling

For sablefish Anoplopoma fimbria that had been transferred abruptly from ambient (5·7° C) to temperatures ranging from 15 to 20° C for 30 min followed by 15 min in air (19·5) C), mortality increased with temperature. Mortality occurred at lower temperatures for sablefish that were net-towed for 4 h at ambient temperature before exposure to a rapid increase in temperature. A clear relationship was apparent between serum lactate and temperature with lactate increasing as temperature increased. For treatments in which mortality did not occur, lactate decreased sharply within 24 h, suggesting recovery. It would appear that the critical postcapture temperature for sablefish that reside and are captured at 4-6° C, would be between 12 and 15° C. The results of this study suggest that fishery management strategies designed to increase postcapture survival of sablefish bycatch should include a consideration of the impact of exposure to seasonal thermoclines and seasonally elevated air temperatures.     

Stem wood properties of mature Norway spruce after 3 years of continuous exposure to elevated [CO2] and temperature

The objective of the study was to investigate the interactive effects of elevated atmospheric carbon dioxide concentration, [CO₂], and temperature on the wood properties of mature field-grown Norway spruce ( Picea abies (L.) Karst.) trees. Material for the study was obtained from an experiment in Flakaliden, northern Sweden, where trees were grown for 3 years in whole-tree chambers at ambient (365 μmol mol⁻¹) or elevated [CO₂] (700 μmol mol⁻¹) and ambient or elevated air temperature (ambient +5.6 °C in winter and ambient +2.8 °C in summer). Elevated temperature affected both wood chemical composition and structure, but had no effect on stem radial growth. Elevated temperature decreased the concentrations of acetone-soluble extractives and soluble sugars, while mean and earlywood (EW) cell wall thickness and wood density were increased. Elevated [CO₂] had no effect on stem wood chemistry or radial growth. In wood structure, elevated [CO₂] decreased EW cell wall thickness and increased tracheid radial diameter in latewood (LW). Some significant interactions between elevated [CO₂] and temperature were found in the anatomical and physical properties of stem wood (e.g. microfibril angle, and LW cell wall thickness and density). Our results show that the wood material properties of mature Norway spruce were altered under exposure to elevated [CO₂] and temperature, although stem radial growth was not affected by the treatments.     

The influence of elevated CO2 and O3 concentrations on Scots pine needles: changes in starch and secondary metabolites over three exposure years

Scots pine (Pinus sylvestris L.) trees, aged about 20 years old, growing on a natural pine heath were exposed to two concentrations of CO₂ (ambient CO₂ and double-ambient CO₂) and two O₃ regimes (ambient O₃ and double-ambient O₃) and their combination in open-top chambers during growing seasons 1994, 1995 and 1996. Concentrations of foliar starch and secondary compounds are reported in this paper. Starch concentrations remained unaffected by elevated CO₂ and/or O₃ concentrations during the first 2 study years. But in the autumn of the last study year, a significantly higher concentration of starch was found in current-year needles of trees exposed to elevated CO₂ compared with ambient air. There were large differences in concentrations of starch and secondary compounds between individual trees. Elevated concentrations of CO₂ and/or O₃ did not have any significant effects on the concentrations of foliar total monoterpenes, total resin acids or total phenolics. Significantly higher concentrations of monoterpenes and resin acids and mostly lower concentrations of starch were found in trees growing without chambers than in those growing in open-top chambers, while there were no differences in concentrations of total phenolics between trees growing without or in chambers. The results suggest that elevated concentrations of CO₂ might increase foliar starch concentrations in Scots pine, while secondary metabolites remain unaffected. Realistically elevated O₃ concentrations do not have clear effects on carbon allocation to starch and secondary compounds even after 3 exposure years. Received: 2 June 1997 / Accepted: 12 December 1997     

Elevated red myotomal muscle temperatures in the most basal tuna species, Allothunnus fallai

The present study tested the hypothesis that Allothunnus fallai can elevate its slow-oxidative red myotomal muscle (RM) temperature. Measurements on 30 A. fallai (750–850 mm fork length) captured by hook and line off the coast of southern New Zealand revealed that RM temperatures are elevated by mean ± s . d . 8·1 ± 1·3° C (range 6·7–10·0° C) above the mean ± s . d . ambient sea surface temperature 15·3 ± 0·8° C (range 14·3 to 16·4° C). These data provide evidence that the vascular modifications to the central circulation of A. fallai act as a counter-current heat exchanger and that RM heat conservation is a character state present in all extant tuna species.     

Synchronization of larval emergence in winter moth (Operophtera brumata L.) and budburst in pedunculate oak (Quercus robur L.) under simulated climate change

1. The hypothesis that a 3 °C elevation in temperature and doubled CO₂ concentration would have no effect on the synchronization of winter moth egg hatch with budburst in oak was tested by comparing the separate and interactive effects of ambient and elevated (+ 3 °C) temperature and ambient and elevated (doubled to 340 p.p.m.) CO₂ in eight experimental Solardomes. In addition, an outdoor control was compared with the ambient temperature/CO₂ treatment combination.
2. Elevated temperature accelerated darkening (preceding egg hatch by about 5–10 days) and hatching of eggs developing off the trees; elevated CO₂ had no effect. The same effects were observed in eggs developing on the trees.
3. Within treatments, date of egg hatch was the same on trees with early or late budburst.
4. Egg darkening and budburst were closely synchronized at both ambient and elevated temperatures.
5. Both eggs and trees required fewer cumulative heat units (day degrees > 4 °C), for hatching and budburst, respectively, at ambient than elevated temperatures. The requirements in the outdoor control treatment were similar to those in the ambient Solardome treatment.
6. Egg hatch between 10 and 25 °C, on a temperature gradient in the laboratory, required a constant number of heat units; fewer were required below 10 °C.
7. Elevated temperatures, in the Solardomes and the field, delayed adult emergence from the pupae.
8. The results suggest that a general increase in temperature with climatic change would not affect the closeness of the synchronization between egg hatch of winter moth and budburst of oak.     

Impacts of elevated CO2 and temperature on soil respiration in warm temperate evergreen Quercus glauca stands: an open-top chamber experiment

We conducted an open-top chamber experiment for 3?years to examine the effect of elevated CO₂ and temperature on soil respiration in experimental stands of Quercus glauca, an evergreen tree species common in the warm temperate zone of Japan. Seedlings of Q. glauca were planted in open-top chambers and treated with factorial combinations of ambient and elevated (ambient?×?1.4, ambient?×?1.8) CO₂ concentrations and ambient and elevated (+3°C) air temperatures. Elevated CO₂ significantly increased the total soil respiration rate (P?<?0.001) and the soil respiration rate at 15°C (R ₁₅) (P?<?0.05) but had no significant effect on the temperature coefficient Q ₁₀. Although temperature significantly affected total soil respiration rate (P?<?0.05), neither the R ₁₅ nor the Q ₁₀ of total soil respiration was affected significantly by the air temperature increase. Annual soil respiration rate, estimated from R ₁₅, Q ₁₀, and soil temperature data, tended to increase with elevated CO₂ concentration. These results suggest that soil respiration rate in Japanese warm temperate broad-leaved forests dominated by Q. glauca is sensitive to elevated CO₂ and is likely to increase under future climatic conditions.     

Lethal effects of experimental warming approximating a future climate scenario on southern African quartz-field succulents: a pilot study

Here we examine the response of succulents in a global biodiversity hot spot to experimental warming consistent with a future African climate scenario. Passive daytime warming (averaging 5.5 degrees C above ambient) of the natural vegetation was achieved with 18 transparent hexagonal open-top chamber arrays randomized in three different quartz-field communities. After 4-months summer treatment, the specialized-dwarf and shrubby succulents displayed between 2.1 and 4.9 times greater plant and canopy mortalities in the open-top chambers than in the control plots. Those surviving in cooler ventilated areas and shaded refuges in the chambers had lower starch concentrations and water contents; the shrubby succulents also exhibited diminished chlorophyll concentrations. It is concluded that current thermal regimes are likely to be closely proximate to tolerable extremes for many endemic succulents in the region, and that anthropogenic warming could significantly exceed their thermal thresholds. Further investigation is required to elucidate the importance of associated moisture deficits in these warming experiments, a potential consequence of supplementary (fog and dew) precipitation interception by open-top chambers and higher evaporation therein, on plant mortalities.     

Influence of simulated warming using OTC on physiological–biochemical characteristics of Elymus nutans in alpine meadow on Qinghai-Tibetan plateau

Elymus nutans Griseb. is a typical important plant species in the alpine meadow of Qinghai-Tibetan plateau. To examine the effects of temperature elevation on its physiological and chemical characteristics, a simulation study was conducted in situ with open-top chambers (OTC) followed the method of International Tundra Experiment (ITEX) from November 2002 to September 2007, and these OTCs were designed five kinds of size with bottom diameters of 0.85, 1.15, 1.45, 1.75, 2.05 m so as to rise different air temperatures. The air temperature inside OTCs increased by 2.68, 1.57, 1.20, 1.07 and 0.69 °C with increase of OTC diameter compared with ambient air. We found that with increase of air temperature, the soluble sugar content and SOD (superoxide dismutase) activity in leaves of E. nutans increased first, and then decreased, whereas, the soluble protein content and GSH (Glutathione) content decreased first and increased then, the chlorophyll a and total chlorophyll contents were decreased, but the contents of chlorophyll b were higher than that of control. Increased temperature enhanced the above-ground biomass and blade height of E. nutans. These results indicated that elevated temperature had significant and complicated effects on physiological–biochemical characteristics of E. nutans on Qinghai-Tibet plateau, when the temperature increased within the range of 0.69–1.57 °C, it may have positive effects on plant growth and development, and E. nutans could adapt even develop defensive strategy to the changes of a certain ecological environment changes.     

Gas exchange and carbon isotope composition of Ananas comosus in response to elevated CO2 and temperature

Ananas comosus L. (Merr.) (pineapple) was grown at three day/night temperatures and 350 (ambient) and 700 (elevated) μ mol mol^–1 CO₂ to examine the interactive effects of these factors on leaf gas exchange and stable carbon isotope discrimination ( Δ ,‰). All data were collected on the youngest mature leaf for 24 h every 6 weeks. CO₂ uptake (mmol m^–2 d^–1) at ambient and elevated CO₂, respectively, were 306 and 352 at 30/20 °C, 175 and 346 at 30/25 °C and 187 and 343 at 35/25 °C. CO₂ enrichment enhanced CO₂ uptake substantially in the day in all environments. Uptake at night at elevated CO₂, relative to that at ambient CO₂, was unchanged at 30/20 °C, but was 80% higher at 30/25 °C and 44% higher at 35/25 °C suggesting that phosphoenolpyruvate carboxylase was not CO₂-saturated at ambient CO₂ levels and a 25 °C night temperature. Photosynthetic water use efficiency (WUE) was higher at elevated than at ambient CO₂. Leaf Δ -values were higher at elevated than at ambient CO₂ due to relatively higher assimilation in the light. Leaf Δ was significantly and linearly related to the fraction of total CO₂ assimilated at night. The data suggest that a simultaneous increase in CO₂ level and temperature associated with global warming would enhance carbon assimilation, increase WUE, and reduce the temperature dependence of CO₂ uptake by A. comosus .     

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.     

Habitat choice in shoals of roach as a function of water temperature and feeding rate

Temperature and food availability are two important factors which affect fish growth and therefore are expected to influence habitat choice in fish. In this study, shoals of 16 juvenile roach, Rutilus rutilus , were given a choice between two chambers that differed in temperature by 1·5°C or 3°C whereas food availability was the same in both chambers (ratio 1 : 1) or higher in the colder one (ratio 4 : 1). The number of fish in each chamber was recorded for 10 min each during a pre-feeding, feeding and post-feeding period. Roach generally preferred the warmer over the colder chamber during the pre-feeding periods. Temperature had a significant effect on the distribution of fish during all three time periods whereas food availability was a significant factor only during the feeding period. The important role of temperature was emphasized further by the fact that a relatively small difference in the temperature gradient of 1·5°C had a stronger effect on fish distribution than a four times higher feeding rate during the feeding period. The implications for growth rates of such short-term decision-making of roach are discussed.     

TEMPERATURE REQUIREMENTS FOR THE DEVELOPMENT AND SURVIVAL OF RICE STEMBORERS IN LABORATORY CONDITIONS

Abstract  The effect of seven constant temperatures from 10 to 40°C (10, 15, 20, 25 30, 35 and 40°C) on the development of eggs, larvae and pupae of rice stemborers viz., Chilo polychrysa (Meyrick), C. suppressalis (Walker), C. partellus (Swinhoe), Scirpophaga incertulas (Walker), S. innotata (Walker) and Sesamia inferens (Walker) were studied. The mean developmental period among constant temperatures (in days) of egg, larva and pupa of six borers differed significantly ( P < 0.0001). The mean percent of development per day of egg, larva and pupa of all borers gradually increased with the increase of constant temperatures. The total developmental period was inversely decreased with the increase of constant temperatures. The lower threshold temperature was found between 10–15°Cand higher threshold temperature between 35–40°C, where no development took place. The mean developmental zero was 8.57±1.71, 7.70±1.01, 8.56±3.25, 10.19±2.19, 8.64±2.68 and 7.91±0.82 for egg, larva and pupa of above-mentioned borers respectively. The total thermal constant of egg, larva and pupa was 705.56, 725.32, 703.30, 556.59, 655.34 and 837.95 degree- days for C. polychrysa, C. suppressalis, C. partellus, S. incertulas, S. innotata and S. inferens respectively. The degree- days required for oviposition of female moths of the six borers was calculated as 99.06, 90.85, 99.29, 75.16, 92.25 and 80.41 respectively. The total degree- days required completing a generation was 804.62, 816.17, 802.59, 631.75, 648.84 and 918.36 respectively.     

The influence of ambient salinity on routine metabolism in the teleost Cyprinodon variegatus Lacepède

Routine metabolism of the euryhaline cyprinodontid Cyprinodon variegatus Lacepede was measured at a series of ambient salinities ranging from fresh water through 100‰. Fish used had been sequentially acclimated to the test salinities (ambient temperature of 20±1° C and a 12: 12 L: D schedule).
Routine metabolic rates were highest at ambient salinities from 15 to 50‰. Metabolism was somewhat lower at ambient salinities less than 15‰, and showed a sequential decline at ambient salinities greater than 50‰. It is suggested that routine metabolism is depressed at elevated salinities by reduced O, transfer, a consequence of maintenance of hydromineral balance in hypersaline waters.     

Responses of the clonal sedge, Carex bigelowii, to two seasons of simulated climate change

The effect of enhanced temperature was studied in open-top chambers on the arctic clonal sedge Carex bigelowii Torr. ex Schwein. at Latnjajaure Field Station, Swedish Lapland for two growing seasons. The flowering phenology of C. bigelowii accelerated during the first field season that the open-top chambers were erected. After the second season of perturbation the sexual reproductive investment was higher in the open-top chambers than in the controls and flowering phenology was still accelerated. Leaf phenology and the growth of vegetative tillers followed the same pattern in the open-top chambers as in the controls, but the leaves were longer in the open-top chambers from the end of June until the end of August. The results suggest that relatively late flowering clonal graminoids may be favoured by warmer climate, at least in the short term.     

Response to simulated climatic change in an alpine and subarctic pollen-risk strategist, Silene acaulis

The aim of this study was to test if early flowering species respond with increased seed production to climate warming as is predicted for late-flowering seed-risk strategists. Experimental climate warming of about 3°C was applied to two populations of the cushion-forming plant Silene acaulis (L.) Jacq. The experiment was run at one subarctic site and one alpine site for 2 years and 1 year, respectively, using open-top chambers (OTC).
The 2-year temperature enhancement at the subarctic site had a marked effect on the flowering phenology. Cushions inside the OTC started flowering substantially earlier than control cushions. Both the male and female phases developed faster in the OTCs, and maturation of capsules occurred earlier. The cushions also responded positively in reproductive terms and produced more mature seeds and had a higher seed/ovule ratio. After 1 year temperature enhancement at the alpine site there was a weak trend for earlier flowering, but there was no significant difference in seed production or seed/ovule ratio.