High-elevation plants have reduced plasticity in flowering time in response to warming compared to low-elevation congeners

Global warming has caused shifts in the flowering time of many plant species. In alpine regions the temperature rise has been especially pronounced and together with decreasing winter precipitation has led to earlier snowmelt. The close association between time of snowmelt and plant growth at high elevations makes climate change for alpine plants particularly threatening. Here we transplanted eleven congeneric pairs of high- and low-elevation herbaceous species to common gardens differing c. 800 m in elevation, and c. 4 °C in mean growing season temperature to test whether reproductive phenologies of high- and low-elevation plants differ in their respective responses to temperature. Results indicate that high-elevation plants were less plastic in response to transplantation than their low-elevation congeners as the onsets of phenophases on average shifted 7 days less than in low-elevation plants. Plasticity of phenophase durations was overall weaker than that of phenophase onsets, and slightly stronger in high-elevation species compared to low-elevation congeners. We suggest that weaker plasticity in the onsets of early stages of reproductive phenology of high-elevation plants is related to spring frost, which constitutes a strong selective agent against early loss of winter hardiness. Some of the plastic responses of both low- and high-elevation species might potentially be adaptive under predicted climate change. However, the observed plasticity can be largely explained as a passive response to temperature and not as the result of natural selection in heterogeneous environments. The strong temperature-sensitivity of low-elevation species might promote their upward range expansion, but only to a certain threshold after which it becomes limited by the short growing season.     

Vegetation development on deposit soils starting at different seasons

Permanent plots were created in different seasons (autumn and spring) and filled with two substrates: nutrient-rich topsoil and nutrient-poor ruderal soil (n = 5 for each treatment). My objectives were to assess the influence of starting season on initial species composition, whether differences at the start cause divergent or convergent pathways of succession and which mechanisms are operating during vegetation development. Mean species richness (number of species per plot) and mean total cover of herb layer differed significantly between substrates and changed significantly during 10 year succession, but there were no significant differences with respect to starting season. However, seasonal as well as substrate effects were evident for particular dominant species and for the pattern of successional sequences. When succession on topsoil plots started in spring, first summer annuals dominated, then monocarpic and polycarpic perennial herbs, then herbaceous perennials together with woody perennials, and at the end of the decade woody perennials. When succession started in autumn, polycarpic perennial herbs dominated from the beginning, and then were replaced by woody perennials in the second half of the decade. On ruderal soil, there was a less rapid but continuous increase of polycarpic perennial herbs and woody species, both on spring and on autumn plots, whereas short-lived plants were more abundant in the first years and then decreased. Species turnover was very high from the first to the second year for all treatments (except topsoil plots starting in autumn), but slowed down during succession. Priority effects due to starting season caused high dissimilarity at the start on the nutrient-rich substrate, but convergent succession towards the end of the first decade. The main mechanisms during early succession on the nutrient-rich topsoil were tolerance based on different life-history traits and inhibition due to reduced light availability. There was no evidence for obligate facilitation. However, an indirect facilitative effect by annuals, which slowed the development of herbaceous perennials down, and thus facilitated growth of woody species, could be seen on topsoil when succession started in spring.     

Molecular control of cold acclimation in trees

Frost tolerance is an acquired characteristic of plants that is induced in response to environmental cues preceding the onset of freezing temperatures and activation of a cold acclimation program. In addition to transient acclimation to low non-freezing temperatures and enhancing survival to short frost episodes during the growth season, perennial woody plants need additionally to survive the cold winter months. Trees have evolved a complex dynamic process controlling the development of dormancy and freezing tolerance that secures accurate initiation and termination of the overwintering process. Although the phenology of overwintering has been known for decades, only recently has there been progress in elucidating the molecular mechanisms of dormancy and freezing tolerance development in perennial plants. Current molecular and genomic studies indicate that herbaceous annual and woody perennial plants share similar cold acclimation mechanisms. Both the signal processes controlling cold acclimation and the cold-regulated target genes appear to be shared by herbaceous and woody plants. However, the dormancy development during overwintering brings new players in the molecular control of seasonal cold acclimation of woody perennials.     

High growing temperatures reduce fruit carbohydrate and vitamin C in kiwifruit

Kiwifruit vines are perennial plants grown in climates varying from maritime to continental. To determine key responses to temperature, vines were heated at different stages of fruit development, and vine growth and fruit composition examined. Heating vines during fruit starch accumulation caused a major shift in partitioning towards vegetative growth and dramatically reduced fruit carbohydrate and vitamin C. In the following season, growth and flowering were severely reduced. Heating vines during fruit cell division had minimal long‐term effects, whereas heating during fruit maturation delayed starch degradation and fruit ripening and affected growth in the following season. When vines were removed from heat, fruit dry matter, starch and sugar levels were always reduced but hexose : sucrose ratios and inositol were raised. Heating vines affected expression of two sucrose synthase genes, but this did not correlate with reduction in fruit carbohydrate. Activity and expression of l ‐galactose dehydrogenase decreased as fruit developed, suggesting some vitamin C biosynthesis must take place in the fruit. Activity and expression of actinidin increased in response to heat. The results of this study have demonstrated both short and long‐term plant responses to elevated temperatures in woody perennials, and that the timing of heat exposure has severe consequences for vitamin C levels in fruit.     

Modulation of relative growth rate and its components by water stress in Mediterranean species with different growth forms

Effects of water availability on seedling growth were analysed in eight Mediterranean species naturally occurring in the Balearic Islands. Seedlings were grown outdoors during summer under two irrigation treatments: field capacity and 35% of field capacity. The relative growth rate (RGR) strongly depended on the growth form, from highest values in herbs to lowest in woody perennials. The main component associated with interspecific variation in RGR was the specific leaf area (SLA), and a quantitative grouping of the different growth forms appeared along the regression line between both parameters. The slow-growing species, i.e. woody perennial shrubs, had the lowest SLA and the fast-growing perennial herbs, the highest, while woody semi-deciduous shrubs appeared intermediate. Decreases in RGR due to water stress were analysed in terms of the relative contribution of the leaf mass ratio (LMR), SLA and the net assimilation rate (NAR). Pooling all species, the decrease in RGR caused by water deficit was mainly explained by decreases in SLA. However, this general pattern was strongly dependent of growth form. Thus, in the woody perennial plants, the decrease in RGR was accompanied by a three-fold decrease in NAR which, however, increased in perennial herbs. SLA increased with decreasing water supply in woody perennial plants, and decreased in woody semi-deciduous shrubs and perennial herbs. Finally, decreases in LMR partly explained decreases in RGR in perennial herbs and woody perennial shrubs. This different response of the different growth forms may reflect differences in seedling adaptation and surviving strategies to drought periods.     

Influence of arbuscular mycorrhiza on growth and reproductive response of plants under water deficit: a meta-analysis

Despite a large body of literature that describes the effects of arbuscular mycorrhizal colonization on plant response to water deficit, reviews of these works have been mainly in narrative form, and it is therefore difficult to quantify the magnitude of the effect. We performed a meta-analysis to examine the effect of mycorrhizal colonization on growth and yield of plants exposed to water deficit stress. Data were compared in the context of annual vs. perennial plants, herbaceous vs. woody plants, field vs. greenhouse conditions, degree of stress, functional group, regions of plant growth, and mycorrhizal and host species. We found that, in terms of biomass measurements, mycorrhizal plants have better growth and reproductive response under water stress compared to non-mycorrhizal plants. When variables such as habit, life cycle, or water stress level are considered, differences in mycorrhizal effect on plant growth between variables are observed. While growth of both annual and perennial plants is improved by symbiosis, perennials respond more favorably to colonization than annuals. Overall, our meta-analysis reveals a quantifiable corroboration of the commonly held view that, under water-deficit conditions, plants colonized by mycorrhizal fungi have better growth and reproductive response than those that are not.     

Differential utilization of summer rains by desert plants

Summary Seasonal changes in the hydrogen isotope ratios of xylem waters were measured to determine water sources used for growth in desert plants of southern Utah. While all species used winter-spring recharge precipitation for spring growth, utilization of summer rains was life-form dependent. Annuals and succulent perennials exhibited a complete dependence on summer precipitation. Herbaceous and woody perennial species simultaneously utilized both summer precipitation and remaining winter-spring precipitation, with herbaceous species much more reliant on the summer precipitation component. Several of the woody perennials exhibited no response to summer precipitation. Currently, precipitation in southern Utah is evenly partitioned between winter and summer time periods; however, global circulation models predict that summer precipitation will increase in response to anticipated climate change. Our data indicate that components within the community will differentially responde to the change in precipitation patterns. These results are discussed in relation to impact on competition and possible changes in community structure.     

Time – size tradeoffs: a phylogenetic comparative study of flowering time, plant height and seed mass in a north-temperate flora

Parents face a timing problem as to when they should begin devoting resources from their own growth and survival to mating and offspring development. Seed mass and number, as well as maternal survival via plant size, are dependent on time for development. The time available in the favorable season will also affect the size of the developing juveniles and their survival through the unfavorable season. Flowering time may thus represent the outcome of such a time partitioning problem. We analyzed correlations between flowering onset time, seed mass, and plant height in a north-temperate flora, using both cross-species comparisons and phylogenetic comparative methods. Among perennial herbs, flowering onset time was negatively correlated with seed mass (i.e. plants with larger seeds started flowering earlier) while flowering onset time was positively correlated with plant height. Neither of these correlations was found among woody plants. Among annual plants, flowering onset time was positively correlated with seed mass. Cross-species and phylogenetically informed analyses largely agreed, except that flowering onset time was also positively correlated with plant height among annuals in the cross-species analysis. The different signs of the correlations between flowering onset time and seed mass (compar. gee regression coefficient=−7.8) and flowering onset time and plant height (compar. gee regression coefficient=+30.5) for perennial herbs, indicate that the duration of the growth season may underlie a tradeoff between maternal size and offspring size in perennial herbs, and we discuss how the partitioning of the season between parents and offspring may explain the association between early flowering and larger seed mass among these plants.     

Host plant predictability and the feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores

Summary Host plant preferences for 34 insect herbivore species are reported. Most polyphagous herbivores feeding on annuals, herbaceous perennials, and woody perennials show distinct preferences for the least abundant plant species among their various host plants. In addition, some populations of widely distributed polyphagous species are much more specialized in their diet than host plant lists alone would suggest. The high level of polyphagy on annuals and herbaceous perennials is suggested to be strongly influenced by the unpredictability of the host plant that is, in turn, controlled by environmental variability. Oligophagous herbivores preferred the least abundant woody perennials on the study sites. Ten of the 22 monophagous herbivores preferred the rarest of all the plant species on the same sites.     

Modelling the Coexistence of Annual and Perennial Plants in Temporally Varying Environments*

Abstract The long-term growth and coexistence of species with large mixed populations in varying environments were modelled for representative environments and life-history characteristics of annual and perennial plants. The effects of the relationships between the means, variances, and covariances of seed yield, establishment, and survival, were explored by Taylor's expansion. The main findings are: 1. Individual variation in reproductive success within generations has no effect on long-term growth, which is determined only by the mean growth rate of the individuals of the species. 2. In annual species with nonoverlapping generations and without seed banks, the species with the largest mean log of the annual growth rate Y, that is the product of the average seed yield per plant and the establishment probability per seed, will win in competition with other species, independent of the correlations between the growth rates of the different species. In this case there is a negative tradeoff between the mean and the variance. 3. In perennial species with a lottery type of equal access to vacant sites, a high annual survival probability allows stable coexistence between perennial species with independent or negatively correlated variance in their mean annual product of seed production and establishment Y. 4. The coexistence range and the likely number of coexisting perennial species increase as a function of the variance of the common species, and is decreased by the variance of the rare species. The coexistence range is decreased by the covariance between the growth rates of the species, and between the survival of the rare species and its growth rate. 5. If mortality in the community of long-lived perennials is synchronized, the generations become nonoverlapping, and the competitive dynamics become similar to that of annuals. 6. Coexistence between annual and perennial species is promoted if the covariances between the annual survival and the relative yield of perennials, and between the yields of perennials and annuals, decrease and become more negative. 7. Selection for seed yield and establishment in different conditions in annuals favours a generalist strategy with low variance between years which provides a moderate yield and establishment over a wide range of environmental conditions. In perennial plants, long-term growth rate is determined by the lifetime seed yield and establishment. Because of strong competition with annuals in the more common conditions, selection in perennials favours instead a specialist strategy, with a high seed yield and establishment at relatively rare occasions in space and time, in which there is only weak competition with annuals. 8. Coexistence of annual species with a long-lived seed bank in the soil is also made possible by independent variation in different years of the germination, seed yield and establishment of different species, analogous to the situation of perennial plants.     

Population dynamics ofHeloniopsis orientalis C. Tanaka (Liliaceae) in natural forests — Sexual reproduction

This study investigated the effects of climate on reproduction, consecutive flowering, and the level of energy investment toward sexual reproduction, as well as the relationships between flowering rates and plant size, and between seed production and plant size. The target species wasHeloniopsis orientalis C. Tanaka (Liliaceae), an evergreen perennial herbaceous plant. Natural populations were monitored in permanent quadrats for seven years in a low-elevation area (Namhansanseong) and for five years in an area with a high elevation (Maranggol). The Tn indices required for flowering were relatively similar each year, but those for seed dispersal fluctuated. The period for seed production was proportional to the total length of the growing season. Rates of flowering varied annually, and depended on conditions in each study area. Flowering was discontinuous (i.e., not consecutive) at Namhansanseong, but was continuous (i.e., consecutive) at Maranggol. The rates were proportional to plant size; the minimum size class that supported flowering was 60–90 cm², and almost all plants with >210 cm² leaf area flowered. The overlapping range in plant size between flowering and non-flowering plants was relatively broad. Numbers of fruit per plant ranged from 1-12, with an average of 6.0 fruits at Namhansanseong and 5.0 fruits at Maranggol. In the latter area, the range in the number of seeds produced and their weight per plant varied drastically, being 1700–1867 seeds and 133.2–177.5 mg, respectively. Almost all correlation coefficients between plant size and fruit number, plant size and seed number, or plant size and seed weight were significant at the 1% level. Lastly, energy investment rates for sexual reproduction were about 30–50% of the net productivity, varying over the years. These rates were proportional to growth rates.     

Temperature response of photosynthesis in C3, C4, and CAM plants: temperature acclimation and temperature adaptation

Most plants show considerable capacity to adjust their photosynthetic characteristics to their growth temperatures (temperature acclimation). The most typical case is a shift in the optimum temperature for photosynthesis, which can maximize the photosynthetic rate at the growth temperature. These plastic adjustments can allow plants to photosynthesize more efficiently at their new growth temperatures. In this review article, we summarize the basic differences in photosynthetic reactions in C₃, C₄, and CAM plants. We review the current understanding of the temperature responses of C₃, C₄, and CAM photosynthesis, and then discuss the underlying physiological and biochemical mechanisms for temperature acclimation of photosynthesis in each photosynthetic type. Finally, we use the published data to evaluate the extent of photosynthetic temperature acclimation in higher plants, and analyze which plant groups (i.e., photosynthetic types and functional types) have a greater inherent ability for photosynthetic acclimation to temperature than others, since there have been reported interspecific variations in this ability. We found that the inherent ability for temperature acclimation of photosynthesis was different: (1) among C₃, C₄, and CAM species; and (2) among functional types within C₃ plants. C₃ plants generally had a greater ability for temperature acclimation of photosynthesis across a broad temperature range, CAM plants acclimated day and night photosynthetic process differentially to temperature, and C₄ plants was adapted to warm environments. Moreover, within C₃ species, evergreen woody plants and perennial herbaceous plants showed greater temperature homeostasis of photosynthesis (i.e., the photosynthetic rate at high-growth temperature divided by that at low-growth temperature was close to 1.0) than deciduous woody plants and annual herbaceous plants, indicating that photosynthetic acclimation would be particularly important in perennial, long-lived species that would experience a rise in growing season temperatures over their lifespan. Interestingly, across growth temperatures, the extent of temperature homeostasis of photosynthesis was maintained irrespective of the extent of the change in the optimum temperature for photosynthesis (T _opt), indicating that some plants achieve greater photosynthesis at the growth temperature by shifting T _opt, whereas others can also achieve greater photosynthesis at the growth temperature by changing the shape of the photosynthesis–temperature curve without shifting T _opt. It is considered that these differences in the inherent stability of temperature acclimation of photosynthesis would be reflected by differences in the limiting steps of photosynthetic rate.     

Do perennials really senesce?

Although senescence remains less studied in perennials than in monocarpic plants, major advances in understanding senescence in perennials have been achieved recently. This success is due not only to the use of genetic tools in woody plants but also to a renaissance of research on how perennials can live for centuries or even millennia. The particularities of perennial life are considered here, with an emphasis on how these affect senescence at different levels of organization. I conclude that although cellular and leaf senescence do share common features in monocarpic and perennial plants, the indeterminacy of meristems found in perennials begs the question of whether senescence really occurs in these organisms at the whole-plant level.     

Establishing woody perennials on hostile soils in arid and semi-arid regions – A review

Background and aims

Woody perennials can be difficult to establish on harsh soils in arid and semi-arid regions. Historically, technological advances have focussed on methods to improve transplanting and direct-seeding but the available information on these advances remains fragmented and the edaphic factors have been largely ignored. This review explores the literature on plant establishment and identifies soil properties that limit plant response in harsh environments.

Conclusions

We reveal that some woody perennials are particularly well-adapted to dry conditions and can also help reclaim degraded landscapes. Furthermore, the environmental and phenological factors that limit the success of direct seeding are well understood but the edaphic factors are not. For example, seedbed preparation and subsoil amelioration before seeding have not been evaluated in dry regions. Seed-priming and seed-placement are also poorly understood, as is the tolerance of woody perennials to different salt types in waterlogged soils of extreme pH and high soil strength. The reason why woody perennials can penetrate strong, hard soils is not obvious from the literature. They apparently cannot exert root growth pressures of the same magnitude as domesticated plants, so they must be able to exploit soil biopores and cracks more efficiently. Other gaps in our understanding of the soil factors that limit woody perennial establishment on hostile soils are identified.     

Age-determination of Dicotyledonous Herbaceous Perennials by Means of Annual Rings: Exception or Rule?

Thirty-five herbaceous dicotyledonous perennial plant species,with permanent root systems, from 16 families, were examinedfor the presence of growth rings in the secondary root xylem.Most of the species surveyed showed ring zonations in the roots,and these could be verified as annual growth rings in the tenspecies for which plants of known age were available. The potentialvalue of ‘herbchronology’ as a tool in ecologicalinvestigations of species and stands of perennial herbs of temperatezones is discussed. Annual rings; age-determination; dendroecology; herbaceous perennials; anatomical patterns; secondary root xylem     

Implications of a temperature increase for host plant range: predictions for a butterfly

Although changes in phenology and species associations are relatively well‐documented responses to global warming, the potential interactions between these phenomena are less well understood. In this study, we investigate the interactions between temperature, phenology (in terms of seasonal timing of larval growth) and host plant use in the polyphagous butterfly Polygonia c‐album. We found that the hierarchy of larval performance on three natural host plants was not modified by a temperature increase as such. However, larval performance on each host plant and temperature treatment was affected by rearing season. Even though larvae performed better at the higher temperature regardless of the time of the rearing, relative differences between host plants changed with the season. For larvae reared late in the season, performance was always better on the herbaceous plant than on the woody plants. In this species, it is likely that a prolonged warming will lead to a shift from univoltinism to bivoltinism. The demonstrated interaction between host plant suitability and season means that such a shift is likely to lead to a shift in selective regime, favoring specialization on the herbaceous host. Based on our result, we suggest that host range evolution in response to temperature increase would in this species be highly contingent on whether the population undergoes a predicted shift from one to two generations. We discuss the effect of global warming on species associations and the outcome of asynchrony in rates of phenological change.     

Long‐term plant community trajectories suggest divergent responses of native and non‐native perennials and annuals to vegetation removal and seeding treatments

Land managers frequently apply vegetation removal and seeding treatments to restore ecosystem function following woody plant encroachment, invasive species spread, and wildfire. However, the long‐term outcome of these treatments is unclear due to a lack of widespread monitoring. We quantified how vegetation removal (via wildfire or management) with or without seeding and environmental conditions related to plant community composition change over time in 491 sites across the intermountain western United States. Most community metrics took over 10 years to reach baseline conditions posttreatment, with the slowest recovery observed for native perennial cover. Total cover was initially higher in sites with seeding after vegetation removal than sites with vegetation removal alone, but increased faster in sites with vegetation removal only. Seeding after vegetation removal was associated with rapidly increasing non‐native perennial cover and decreasing non‐native annual cover. Native perennial cover increased in vegetation removal sites irrespective of seeding and was suppressed by increasing non‐native perennial cover. Seeding was associated with higher non‐native richness across the monitoring period as well as initially higher, then declining, total and native species richness. Several cover and richness recovery metrics were positively associated with mean annual precipitation and negatively associated with mean annual temperature, whereas relationships with weather extremes depended on the lag time and season. Our results suggest that key plant groups, such as native perennials and non‐native annuals, respond to restoration treatments at divergent timescales and with different sensitivities to climate and weather variation.     

Sulfur Metabolism in Plants: Are Trees Different?

Sulfur metabolite levels and sulfur metabolism have been studied in a significant number of herbaceous and woody plant species. However, only a limited number of datasets are comparable and can be used to identify similarities and differences between these two groups of plants. From these data, it appears that large differences in sulfur metabolite levels, as well as the genetic organization of sulfate assimilation and metabolism do not exist between herbaceous plants and trees. The general response of sulfur metabolism to internal and/or external stimuli, such as oxidative stress, seems to be conserved between the two groups of plants. Thus, it can be expected that, generally, the molecular mechanisms of regulation of sulfur metabolism will also be similar. However, significant differences have been found in fine tuning of the regulation of sulfur metabolism and in developmental regulation of sulfur metabolite levels. It seems that the homeostasis of sulfur metabolism in trees is more robust than in herbaceous plants and a greater change in conditions is necessary to initiate a response in trees. This view is consistent with the requirement for highly flexible defence strategies in woody plant species as a consequence of longevity. In addition, seasonal growth of perennial plants exerts changes in sulfur metabolite levels and regulation that currently are not understood. In this review, similarities and differences in sulfur metabolite levels, sulfur assimilation and its regulation are characterized and future areas of research are identified.     

Variation in the establishment of a non-native annual grass influences competitive interactions with Mojave Desert perennials

Competition between native and non-native species can change the composition and structure of plant communities, but in deserts, the highly variable timing of resource availability also influences non-native plant establishment, thus modulating their impacts on native species. In a field experiment, we varied densities of the non-native annual grass Bromus madritensis ssp. rubens around individuals of three native Mojave Desert perennials—Larrea tridentata, Achnatherum hymenoides, and Pleuraphis rigida—in either winter or spring. For comparison, additional plots were prepared for the same perennial species and seasons, but with a mixture of native annual species as neighbors. Growth of perennials declined when Bromus was established in winter because Bromus stands had 2–3 months of growth and high water use before perennial growth began. However, water potentials for the perennials were not significantly reduced, suggesting that direct competition for water may not be the major mechanism driving reduced perennial growth. The impact of Bromus on Larrea was lower than for the two perennial grasses, likely because Larrea maintains low growth rates throughout the year, even after Bromus has completed its life cycle. This result contrasts with the perennial grasses, whose phenology completely overlaps with (Achnatherum) or closely follows (Pleuraphis) that of Bromus. In comparison, Bromus plants established in spring were smaller than those established in winter and thus did not effectively reduce growth of the perennials. Growth of perennials with mixed annuals as neighbors also did not differ from those with Bromus neighbors of equivalent biomass, but stands of these native annuals did not achieve the high biomass of Bromus stands that were necessary to reduce perennial growth. Seed dormancy and narrow requirements for seedling survivorship of native annuals produce densities and biomass lower than those achieved by Bromus; thus, impacts of native Mojave Desert annuals on perennials are expected to be lower than those of Bromus.