Germination at extreme temperatures : implications for alpine shrub encroachment

Worldwide, shrub cover is increasing across alpine and tundra landscapes in response to warming ambient temperatures and declines in snowpack. With a changing climate, shrub encroachment may rely on recruitment from seed occurring outside of the optimum temperature range. We used a temperature gradient plate in order to determine the germination niche of 14 alpine shrub species. We then related the range in laboratory germination temperatures of each species to long-term average temperature conditions at: (1) the location of the seed accession site and (2) across each species geographic distribution. Seven of the species failed to germinate sufficiently to be included in the analyses. For the other species, the germination niche was broad, spanning a range in temperatures of up to 17 ◦C, despite very low germination rates in some species. Temperatures associated with the highest germination percentages were all above the range of temperatures present at each specific seed accession site. Optimum germination temperatures were consistently within or higher than the range of maximum temperatures modelled across the species’ geographic distribution. Our results indicate that while some shrub species germinate well at high temperatures, others are apparently constrained by an inherent seed dormancy. Shrub encroachment in alpine areas will likely depend on conditions that affect seed germination at the microsite-scale, despite overall conditions becoming more suitable for shrubs at high elevations

Effects of warming temperatures on germination responses and trade-offs between seed traits in an alpine plant

1. Climate warming may affect multiple aspects of plant life history, including important factors such as germination responses and the key trade-off between offspring size and number. As a case study to address these concepts, we used an alpine plant (waxy bluebell, Wahlenbergia ceracea; Campanulaceae) that shows plasticity to warming in seed traits and in which seed dormancy status regulates germination. We chose an alpine species because alpine environments are ecosystems particularly under threat by climate change. 2. We conducted germination assays under cool and warm temperatures using seeds produced by individuals that were grown under historical (cooler) and future (warmer) temperature scenarios. We assessed the presence of a seed size vs number trade-off, and then examined the effects of seed number and size on germination percentage, the fractions of dormant and viable seeds, and germination velocity. Further, we examined whether warming during parental growth and during germination affected these relationships. 3. We found evidence for a seed size vs number trade-off only under historical parental temperatures. Indeed, under future growth temperatures, parental plants produced fewer and smaller seeds and there was no evidence of a trade-off. However, the reductions in both seed traits under warming did not affect germination, despite correlations of seed size and number with germination traits. Warming increased germination, particularly of larger seeds, but overall it resulted in more than fourfold reductions in parental fitness. 4. Synthesis. Our study shows the importance of growth conditions when evaluating the seed size vs number trade-off. Stressful conditions, such as warmer temperatures, can restrain the ability of plants to reach optimal investment in reproduction, masking the trade-off. By analysing responses across the whole life cycle, we show here an overall detrimental effect of warming, highlighting the potential risk of climate change for W. ceracea, and, potentially, for alpine plant communities more widely.Files can be opened using Excel and analysed using R.Funding provided by: Australian Research CouncilCrossref Funder Registry ID: http://dx.doi.org/10.13039/501100000923Award Number: DP170101681Experiments were conducted using the plant species Wahlnebrgia ceracea (waxy bluebells). Two datasets were used in this manuscript. 1) Seed size vs number trade-off: Parental individuals from a total of 30 lines ('Line') were grown in growth chambers for 191 days under temperature conditions of a historical/cooler (1960–1970) or a projected future/warmer (2090–2100) climate ('Parental_Temperature'). The parental individuals were randomly assigned to one of three blocks, which corresponded to positions inside the chambers, and each block was equivalent in all chambers ('Block'). Day and night temperatures during the experiment were changed every 15 days to mimic seasonality, with the maximum day temperatures during the peak of summer being 24°C and 29°C for the historical and future parental temperatures, respectively. After 100 days since the imposition of the temperature treatments (during the peak of the summer), half of the plants were moved for 5 days to new chambers, where the temperature was 5°C above the respective treatments, i.e., 29°C and 34°C ('Heatwave'). After this time, the parental individuals were moved back to their respective historical or future temperature treatments. We collected the seeds throughout the 191 days of parental growth, and we stored them in desiccators for at least 11 weeks. After this time, we calculated seed size ('Seed_Size') as the average mass of three lots of 50 seeds divided by 50. We calculated seed number ('Seed_Number') as the ratio between the cumulative mass of the seeds produced by each parental individual and seed size. The 30 lines of the parental individuals were obtained by crossing plants that originated from seeds that were collected at the same elevation, either high or low elevation ('Elevation') in sites within Kosciuszko National Park, NSW, Australia. Therefore, 14 lines originated from high elevations and 14 lines from low elevations. 2) Germination responses - seed traits correlations: The seeds were harvested from the parental individuals grown under historical/cooler or projected future/warmer temperatures ('Parental_Temperature') (see above) from a subset of 14 lines ('Line'). These seeds were used in germination assays in the glasshouse under cool (25°C) or warm temperatures (30°C) ('Germination_Temperature'). We measured seed size ('Seed_Size') as the average mass of three lots of 50 seeds; then these seeds were sowed in agar dishes (25 seeds per dish, 2 dishes per temperature treatment from each parental individual). Seed number ('Seed_Number') was the same as above. Dishes were left under temperature treatments for 4 weeks to allow germination of the non-dormant fraction of the seeds ('Not_Dormant_Seeds') and germination was checked once per week. Then, all the dishes were moved to a cold room at 4–5°C in the dark for 4 weeks to allow cold stratification. After this time, dishes were moved back to the glasshouse under the same temperature treatments as before to allow germination of the dormant seeds. We considered seeds to be dormant ('Dormant_seeds') if they germinated during or after cold stratification or if they did not germinate at all but were still determined to be viable at the end of the experiment. We considered seed to be viable ('Viable_Seeds') if they germinated ('Germinated_Seeds') as well as the seeds that contained an endosperm but still did not germinate ('Not_Germinated_Seeds'), while we considered empty seeds as non-viable ('Not_Viable_Seeds'). Germinated and not germinated seeds (as above) were used to calculate the germination percentage. We calculated germination velocity ('Germination_Velocity') as the reciprocal of the mean germination time (germination velocity (germination (%) week-1) GV = (G1 + G2 +…+ Gn) / (G1 x T1 + G2 x T2 +…+ Gn x Tn), where Gn is the number of new germinating seeds at each sampling point, and Tn is the time between each sampling point (= one week). The files provided present the datasets in their first sheet and keys with the definitions of each term in the second sheet

Tolerance of warmer temperatures does not confer resilience to heatwaves in an Alpine herb

Climate change is generating both sustained trends in average temperatures and higher frequency and intensity of extreme events. This poses a serious threat to biodiversity, especially in vulnerable environments, like alpine systems. Phenotypic plasticity is considered to be an adaptive mechanism to cope with climate change in situ, yet studies of the plastic responses of alpine plants to high temperature stress are scarce. Future weather extremes will occur against a background of warmer temperatures, but we do not know whether acclimation to warmer average temperatures confers tolerance to extreme heatwaves. Nor do we know whether populations on an elevational gradient differ in their tolerance or plasticity in response to warming and heatwave events. We investigated the responses of a suite of functional traits of an endemic Australian alpine herb, Wahlenbergia ceracea, to combinations of predicted future (warmer) temperatures and (relative) heatwaves. We also tested whether responses differed between high- vs. low-elevation populations. When grown under warmer temperatures, W. ceracea plants showed signs of acclimation by means of higher thermal tolerance (Tcrit, T50, and Tmax). They also invested more in flower production, despite showing a concurrent reduction in photosynthetic efficiency (Fv/Fm) and suppression of seed production. Heatwaves reduced both photosynthetic efficiency and longevity. However, we found no evidence that acclimation to warmer temperatures conferred tolerance of the photosynthetic machinery to heatwaves. Instead, when exposed to heatwaves following warmer growth temperatures, plants had lower photosynthetic efficiency and underwent a severe reduction in seed production. High- and low-elevation populations and families exhibited limited genetic variation in trait means and plasticity in response to temperature. We conclude that W. ceracea shows some capacity to acclimate to warming conditions but there is no evidence that tolerance of warmer temperatures confers any resilience to heatwaves.This research was supported by the Australian Research Council (DP170101681), an International Ph.D. Scholarship to RN and an ARC Future Fellowship FT110100453 to LK. Research grants funded all research related costs (such as renting growth chambers or buying equipment), while the scholarship paid a stipend to RN

A high-throughput method for measuring critical thermal limits of leaves by chlorophyll imaging fluorescence

Plant thermal tolerance is a crucial research area as the climate warms and extreme weather events become more frequent. Leaves exposed to temperature extremes have inhibited photosynthesis and will accumulate damage to PSII if tolerance thresholds are exceeded. Temperature-dependent changes in basal chlorophyll fluorescence (T-F0) can be used to identify the critical temperature at which PSII is inhibited. We developed and tested a high-throughput method for measuring the critical temperatures for PSII at low (CTMIN) and high (CTMAX) temperatures using a Maxi-Imaging fluorimeter and a thermoelectric Peltier plate heating/cooling system. We examined how experimental conditions of wet vs dry surfaces for leaves and heating/cooling rate, affect CTMIN and CTMAX across four species. CTMAX estimates were not different whether measured on wet or dry surfaces, but leaves were apparently less cold tolerant when on wet surfaces. Heating/cooling rate had a strong effect on both CTMAX and CTMIN that was species-specific. We discuss potential mechanisms for these results and recommend settings for researchers to use when measuring T-F0. The approach that we demonstrated here allows the high-throughput measurement of a valuable ecophysiological parameter that estimates the critical temperature thresholds of leaf photosynthetic performance in response to thermal extremes.This research was supported by the Australian Research Council (DP170101681)

An Automatic Palindrome Generator

In 1984 Dan Hoey, a US naval mathematician, wrote a computer which he used to create a 540-word expansion of Leigh Mercer\u27s Panama palindrome (PD). It began A man, a plan, a caret, a ban, a myriad, a sum, a lac... and ended ...a calmus, a diaryman, a bater, a canal Panama. (For the full PD, plus additional information, see http://www2.vo.lu/homepages/phahn/anagrams/panama/htm.

Predicting species and community responses to global change using structured expert judgement : an Australian mountain ecosystems case study

Conservation managers are under increasing pressure to make decisions about the allocation of finite resources to protect biodiversity under a changing climate. However, the impacts of climate and global change drivers on species are outpacing our capacity to collect the empirical data necessary to inform these decisions. This is particularly the case in the Australian Alps which has already undergone recent changes in climate and experienced more frequent large-scale bushfires. In lieu of empirical data, we used a structured expert elicitation method (the IDEA protocol) to estimate the abundance and distribution of nine vegetation groups and 89 Australian alpine and subalpine species by the year 2050. Experts predicted that most alpine vegetation communities would decline in extent by 2050; only woodlands and heathlands are predicted to increase in extent. Predicted species-level responses for alpine plants and animals were highly variable and uncertain. In general, alpine plants spanned the range of possible responses, with some expected to increase, decrease or not change in cover. By contrast, almost all animal species are predicted to decline or not change in abundance or elevation range; more species with water-centric life-cycles are expected to decline in abundance than other species. While long-term ecological data will always be the gold-standard in informing the future of biodiversity, the method and outcomes outlined here provide a pragmatic and coherent basis upon which to start informing conservation policy and management in the face of rapid change and paucity of data

Landscape genomic prediction for restoration of a Eucalyptus foundation species under climate change

As species face rapid environmental change, we can build resilient populations through restoration projects that incorporate predicted future climates into seed sourcing decisions. Eucalyptus melliodora is a foundation species of a critically endangered community in Australia that is a target for restoration. We examined genomic and phenotypic variation to make empirical based recommendations for seed sourcing. We examined isolation by distance and isolation by environment, determining high levels of gene flow extending for 500 km and correlations with climate and soil variables. Growth experiments revealed extensive phenotypic variation both within and among sampling sites, but no site-specific differentiation in phenotypic plasticity. Model predictions suggest that seed can be sourced broadly across the landscape, providing ample diversity for adaptation to environmental change. Application of our landscape genomic model to E. melliodora restoration projects can identify genomic variation suitable for predicted future climates, thereby increasing the long term probability of successful restoration

The thermal tolerance of photosynthetic tissues: a global systematic review and agenda for future research

Understanding plant thermal tolerance is fundamental to predicting impacts of extreme temperature events that are increasing in frequency and intensity across the globe. Extremes, not averages, drive species evolution, determine survival, and increased crop performance. To better prioritise agricultural and natural system research, it is crucial to evaluate how researchers are assessing the capacity of plants to tolerate extreme events. We conducted a systematic review to determine how plant thermal tolerance research is distributed across wild and domesticated plants, growth forms and biomes, and identify crucial knowledge gaps. Our review shows that most thermal tolerance research examines cold tolerance of cultivated species; ~5% of articles consider both heat and cold tolerance. Plants of extreme environments are understudied, and techniques widely applied in cultivated systems are largely unused in natural systems. Lastly, we find that lack of standardised methods and metrics compromises the potential for mechanistic insight. Our review provides an entry point for those new to the methods used in plant thermal tolerance research and bridges often disparate ecological and agricultural perspectives for the more experienced. We present a considered agenda of thermal tolerance research priorities to stimulate efficient, reliable, and repeatable research across the spectrum of plant thermal tolerance

Reframing conservation physiology to be more inclusive, integrative, relevant and forward-looking: reflections and a horizon scan

Applying physiological tools, knowledge and concepts to understand conservation problems (i.e. conservation physiology) has become common place and confers an ability to understand mechanistic processes, develop predictive models and identify cause-and-effect relationships. Conservation physiology is making contributions to conservation solutions; the number of 'success stories' is growing, but there remain unexplored opportunities for which conservation physiology shows immense promise and has the potential to contribute to major advances in protecting and restoring biodiversity. Here, we consider howconservation physiology has evolved with a focus on reframing the discipline to be more inclusive and integrative. Using a 'horizon scan', we further exploreways in which conservation physiology can be more relevant to pressing conservation issues of today (e.g. addressing the Sustainable Development Goals; delivering science to support the UN Decade on Ecosystem Restoration), as well as more forward-looking to inform emerging issues and policies for tomorrow. Our horizon scan provides evidence that, as the discipline of conservation physiology continues to mature, it provides a wealth of opportunities to promote integration, inclusivity and forward-thinking goals that contribute to achieving conservation gains. To advance environmenta lmanagement and ecosystem restoration, we need to ensure that the underlying science (such as that generated by conservation physiology) is relevant with accompanying messaging that is straightforward and accessible to end users