Phenological responses to nitrogen and water addition are linked to plant growth patterns in a desert herbaceous community

Increases in nitrogen (N) deposition and variation in precipitation have been occurring in temperate deserts; however, little information is available regarding plant phenological responses to environmental cues and their relationships with plant growth pattern in desert ecosystems. In this study, plant phenology and growth of six annuals in response to N and water addition were monitored throughout two consecutive growing seasons in 2011 and 2012 in a temperate desert in northwestern China. The effects of N and water addition on reproductive phenology differed among plant species. N and water addition consistently advanced the flowering onset time and fruiting time of four spring ephemerals; however, their effects on two spring‐summer annuals were inconsistent, with advances being noted in one species and delays in another. N and water addition alone increased plant height, relative growth rate, leaf number, flower number, and individual biomass, while their combinative effects on plant growth and reproductive phenology were dependent on species. Multiple regression analysis showed that flowering onset time was negatively correlated with relative growth rate of two species, and negatively correlated with maximum plant height of the other four species. Our study demonstrates that phenological responses to increasing precipitation and N deposition varied in annuals with different life histories, whereby the effects of climate change on plant growth rate were related to reproductive phenology. Desert annuals that were able to accelerate growth rate under increasing soil resource availability tended to advance their flowering onset time to escape drought later in the growing season. This study promotes our understanding of the responses of temperate desert annuals to increasing precipitation and N deposition in this desert.     

Accumulation of buried seeds and establishment of ruderal therophytic communities in disturbed habitat,central Japan

The establishment and maintenance mechanisms of pioneer communities were investigated in ruderal habitats under two disturbance regimes, frequent and infrequent cutting sites. In the infrequent cutting sites, large perennials dominated through the year and inhibited the invasion of annuals, and the perennial community succeeded to forest stage if the cutting was stopped. In the frequent cutting sites, therophytic communities of winter and summer annuals alternated by season. Fresh seeds of both winter and summer annuals are dormant, but they have different germination times and thus can share the same sites in different seasons. Wind-dispersed biennials and large perennials have nondormant seeds and easily invade the sites; however, they are unable to mature to reproductive phase due to recurrent cuttings. The therophytic species, which can complete their life-cycle in a period between cuttings, accumulate seeds in the soils and are maintained by these buried seeds (seed bank annuals) during recurrent disturbances. The seed bank is compensation for the dispersal inefficiency of seed bank annuals. After abandonment of the frequent cutting sites, the buried seeds of seed bank annuals germinate and become the first-year pioneers. Thus, seed bank annuals are not invasive colonizers but are the remnants of the ruderal weed communities before abandonment.     

Long‐term response of a Mojave Desert winter annual plant community to a whole‐ecosystem atmospheric CO2 manipulation (FACE)

Desert annuals are a critically important component of desert communities and may be particularly responsive to increasing atmospheric (CO₂) because of their high potential growth rates and flexible phenology. During the 10‐year life of the Nevada Desert FACE (free‐air CO₂ enrichment) Facility, we evaluated the productivity, reproductive allocation, and community structure of annuals in response to long‐term elevated (CO₂) exposure. The dominant forb and grass species exhibited accelerated phenology, increased size, and higher reproduction at elevated (CO₂) in a wet El Niño year near the beginning of the experiment. However, a multiyear dry cycle resulted in no increases in productivity or reproductive allocation for the remainder of the experiment. At the community level, early indications of increased dominance of the invasive Bromus rubens at elevated (CO₂) gave way to an absence of Bromus in the community during a drought cycle, with a resurgence late in the experiment in response to higher rainfall and a corresponding high density of Bromus in a final soil seed bank analysis, particularly at elevated (CO₂). This long‐term experiment resulted in two primary conclusions: (i) elevated (CO₂) does not increase productivity of annuals in most years; and (ii) relative stimulation of invasive grasses will likely depend on future precipitation, with a wetter climate favoring invasive grasses but currently predicted greater aridity favoring native dicots.     

PATTERNS OF RESOURCE ALLOCATION IN WILDFLOWER POPULATIONS OF FIELDS AND WOODS

Fifty-two populations of fifty species of wildflowers characteristic of either field or deciduous forest were analyzed to determine pattens of biomass allocation to component organs. These populations' allocation patterns were used to determine generalized allocation patterns of the herb component of earlier (field) and later (deciduous forest) secondary successional communities. The following patterns were determined: (1) The proportion of dry matter allocated to seed reproductive organs was greater in field populations than in woods populations; (2) The herbs of woodland habitats allocated a greater proportion of their resources to leaves and belowground organs than field habitat herbs; (3) The field annuals allocated a greater proportion of their resources to reproduction than field perennials; (4) Introduced species showed a higher reproductive allocation than native species of fields; (5) Regression analysis showed strong correlations of component organ biomass to total biomass and belowground biomass to shoot biomass.     

The ecological significance of annuals on a Mediterranean grassland (Mt Ventoux,France)

A study was carried out on a Mediterranean limestone grassland to search for the ecological significance of the occurrence of annuals. In the studied grassland, most of annuals are concentrated in patches. Authors tested if annuals (1) correspond to the R-strategy (ruderal) according to Grime's theory, i. e. they are adapted to live in disturbed, productive habitats, or (2) if they present an adaptation to a xeric climate in keeping with Raunkiaer.The correspondence analyses and the examination of plants strategies reveal that:(1) annuals are grouped with the more xerophytic perennials in the ordination,(2) the cover of annuals is higher in plots with a high cover of S species (stress-tolerators) and a lower cover of C species (competitors) and CS species (intermediate strategy).These results constitute arguments to integrate annuals of Mediterranean grasslands into stress-tolerators. In the discussion are examined the importance of patches in which annuals are preferentially located. Although patches are generated by disturbances, the presence of annuals is explained by their life cycle fitting the microclimate variations, characterized by an extreme summer drought. Their cycle is not adapted here to the frequency of disturbances (unless the summer drought is considered as annual disturbance - this last point of view is discussed). In the studied grassland, annuals are then better thought of as highly developed stress-tolerators than as ruderals. A generalization is put forward by distinguishing two cases: in xeric habitats, annuals are stress-tolerant, and in productive habitats, they are ruderal.Nomenclature: follows Kerguélen (1993)     

The evolution of self-fertilization in perennials

Abstract Many plants are perennials, but studies of self-fertilization do not usually include features of perennial life histories. We therefore develop models that include selfing, a simple form of perenniality, adult inbreeding depression, and an adult survivorship cost to seed production. Our analysis shows that inbreeding depression in adults diminishes the genetic transmission advantage associated with selfing, especially in long-lived perennials that experience inbreeding depression over many seasons. Perennials also pay a cost when selfing increases total seed set at the expense of future survivorship and reproduction. Such life-history considerations shed new light on the generalization that annuals self-fertilize more than perennials. Past research suggested reproductive assurance as an explanation for this association, but common modes of selfing offer equal reproductive assurance to annuals and perennials. Instead, perennials may avoid selfing because of adult inbreeding depression and the cost to future survivorship and reproduction.     

ROLE OF RECRUITMENT IN STRUCTURING BEDS OF SARGASSUM SPP. (PHAEOPHYTA) AT ROTTNEST ISLAND,WESTERN AUSTRALIA1

The importance of annual recruitment to the structure of adult stands of Sargassum was determined for a mixed species Sargassum bed at Rottnest Island, Western Australia. The morphologically similar species Sargassum spinuligerum Sonder, S. distichum Sonder, and S. podacanthum Sonder grew together in the shallow subtidal (6 m). Positive species determinations were only possible when thalli were reproductive, so recruits, bases, and vegetative annuals for all species were grouped together. Densities of recruits, perennial bases, vegetative annuals, and reproductive annuals were determined at monthly intervals from 20 randomly placed 0.25-m^?2 quadrats. Recruitment and mortality for recruits and adults were further determined at three monthly intervals from 6-×-1-m^?2 permanent quadrats. The density of adults varied little with season (between 32 and 58 m^?2). Growth of annuals was initiated in April, thalli became reproductive by late August–early September, and senescence occurred in December–January. Density of recruits was highly variable (1.6–210 individuals-m^?2) and peaked seasonally during late summer (January–February) and then declined rapidly. Adults showed a complete turnover of thalli in the bed over 25–27 months. Adult mortality was compensated by annual recruitment from propagules (43%) and vegetative regeneration from fragments of holdfasts left on the reef (57%). A seasonal pattern in survivorship was observed for adults that grew from recruits with higher initial numbers and lower mortalities for August and November cohorts. Little seasonally was observed in survivorship of adults that grew vegetatively from remnant crusts. Although initial cohort sizes were smaller for adults grown from recruits than from remnant crusts, mortality was lower, resulting in similar contributions to adult density from both recruits and remnant crusts. Recruitment from propagules and vegetative regeneration played an important role in buffering the adult stand from high rates of mortality and reducing seasonal variation in adult density and contributed to the persistence and seasonal structure of Sargassum beds at Rottnest Island.     

Habitat gradient and reproductive habits of the sevenStellaria species in Japan

The author studied the habitat preference, life form, life cycle and reproductive characteristics of the seven species of the genusStellaria along a valley in the northeastern part of Kyoto City. Along the valley 150 quadrats were set from the village up to the hills. The quadrats were sorted into 15 quadrat groups by their affinity in the environment. The quadrat groups were ordinated by the quotients of community and the Bray-Curtis method in order to reveal the complex gradient in the environment. The complex gradient could be best understood by the degree of human disturbance. The seven species showed a continuous distribution along the complex gradient. There existed three patterns in their habitat preference and reproductive characteristics: (1)S. media andS. neglecta grew as annuals in the habitat of severe human disturbance. (2)S. tomentella, S. sessiliflora, andS. diversiflora grew as perennials in the habitat of slight human disturbance. (3)S. aquatica andS. alsine grew as facultative annuals either in the habitat of severe or slight human disturbance.     

Reproductive phenology and biology in annual and perennial Alismataceae

Two annual (Echinodorus rostratus (Nutt.) Engelm., Sagittaria calycina Engelm.) and 3 perennial (Alisma subcordatum Rat., S. brevirostra Mack & Bush, S. latifolia Wild.) species of Alismataceae were compared to determine differences in reproductive effort and timing. The annuals produce flowers and fruits earlier and later than the perennials and, under optimal conditions, the annuals also produce more (but smaller) seeds annually per plant. Populations of the annuals occur on exposed mud and are erratically abundant from year to year, while populations of the perennials are more stable. Seed viability of the annuals decreases more slowly over time. There is general congruence of these life-history attributes with the concepts of “r” and “K” strategies.     

Background processes at the population level during succession in grasslands on sand

Two population characteristics, reproductive allocation (RA) and the intensity of photosynthesis, were investigated in the successional stages of the sandy grassland communities of the Great Hungarian Plain. Most of the species studied changed their allocational response as succession advanced. Compound path schemes, as casual models, were constructed and analysed for describing intra-individual effects (influence of assimilating organs and root mass of the plant sampled) as well as the phytosociological effects on RA. In pioneer stages, the high values of path coefficients show that the effect of assimilating organs is responsible for the variability of RA. It is also shown that annuals and perennials exhibit different behaviour as to the regulation of RA. It seems that-in terms of path analysis-the reproduction of annuals is influenced by internal factors only. In case of perennials, an external (namely phytosociological) regulation was also observed. It is suggested that successional stages play a role in regulating the photosynthetic intensity of both the dominant species and the whole plant assemblage.Abbreviations RA = Reproductive Allocation     

Analysis of environmental and species effects on the magnitude of biomass investment in the reproductive effort of annual pasture plants

Summary In annual pastures utilized for grazing, the amount and quality of the standing dry matter in the dry season is of importance for the performance of the animals. Often both characteristics decline at the end of the green season. It is shown, that dispersal of the reproductive structures of the vegetation may be the main reason for this phenomenon. Determination of the reproductive effort of the annuals indicates that the proportion of their total production invested in reproductive tissue may be as high as that of cultivated species. It is shown that the harvest index (or seed ratio) of annual species is closely related to nutrient (mainly nitrogen) transfer from vegetative organs to the reproductive organs in the period between flowering and maturity, when in most cases, additional uptake of nitrogen from the soil is negligible. The effect of environmental and genetic effects on these processes is discussed.Present address: Laboratory of Soils and Fertilizers, Agricultural University, P.O. Box 14, Wageningen, The Netherlands     

Plants Growing on the Walls of Italian Towns 2. Reproductive Ecology

Abstract

In this study of the reproductive ecology of wall plants, adaptation to wall habitats was observed to be influenced by characteristics of the vegetative and reproductive structures. Perennial species were more numerous than annuals. Diaspore dispersal was principally anemochorous, due to the small size and weight of diaspores, together with anatomical features and aerostatic organs favouring wind dispersal. Species adaptability and the efficiency of reproductive mechanisms are evidently the main factors ensuring the survival and dispersal of wall plants.     

Perennial polypores as indicators of annual and red-listed polypores

Many polypores are specialized in their requirements for substrate and environment, and they have been suggested to indicate the continuity of coarse woody debris or naturalness of a forest stand. However, the use of polypores as indicators of conservation value is restricted by the temporally limited appearance of annual fruit bodies. We studied whether the species richness of perennial polypores (perennials) can be used to predict the species richness of annual or annual red-listed polypores (annuals). Our data included 1471 separate datasets (sample plots or larger inventoried areas) in different parts of Finland and Russian Karelia, ranging from the southern to northern boreal zone. At the large scale (the whole area) the number of perennials explained about 70% of the variation in the number of annuals, and about 67% in the number of red-listed annuals. A minimum set of 40–60 perennial occurrences gave a reliable estimate on the species richness of annuals, and 60–80 occurrences on the species richness of red-listed annuals. The richness of perennials predicted the richness of annuals and, in particular, richness of red-listed annuals, better than the size of inventoried area. According to our results, perennial polypores can be used as a surrogate for overall polypore species richness in natural and seminatural boreal forests, but the predictive power is weaker in managed forests. In addition, the relationship between the perennial and annual species seems to differ in different vegetation zones, management types and forest types. Due to this variation direct application of the indicator values derived from different vegetation zones and management or forest types are not recommended. Since perennials are easier to identify than annuals, detectable throughout the year, and have much smaller year-to-year variation, their use as an indicator group seems to offer advantages regarding the timing and cost-efficiency of inventories.     

Genetics of flower initiation and development in annual and perennial plants

Flowering is an integral developmental process in angiosperms, crucial to reproductive success and continuity of the species through time. Some angiosperms complete their life cycle within a year (annual plants), and others have a longer reproductive life, which is characterized by the generation of new flowering and vegetative shoots every year (perennial plants). Despite the differences in their lifespan, the underlying genetics of flower induction and floral organ formation appears to be similar among these plants. Hence, the knowledge gained from the study of flowering mechanism in Arabidopsis thaliana can be used to better understand similar processes in other plant species, especially the perennials, which usually have a long generation time and are not amenable to genetic analysis. Using Arabidopsis as a model, we briefly discuss the current understanding of the transition from vegetative to reproductive growth and the subsequent formation of individual floral organs, and how this knowledge has been successfully applied to the identification of homologous genes from perennial crops. Although annuals appear to share many similarities with perennials in terms of gene function, they differ in their commitment to flowering. Once an annual reaches the reproductive phase, all meristems are typically converted into either floral or inflorescence meristems. In contrast, each year, each meristem of a mature perennial has the choice to produce either a vegetative or a reproductive shoot. The physiology and genetics of flowering in Citrus are used to highlight the complexity of reproductive development in perennials, and to discus possible future research directions.     

Photosynthetic capacity in relation to leaf position in desert versus old-field annuals

Summary Desert annuals of Death Valley, California have higher average light-saturated photosynthetic capacities and leaf nitrogen contents than do early-successional annuals of Illinois. The leaves of annuals in the light-unlimited Death Valley environment change little in specific weight, nitrogen, or photosynthetic capacity with age. In contrast, these properties decrease markedly with age in the leaves of the Illinois annuals even in leaves not exposed to the usual shading that accompanies canopy development. These results are interpreted in a carbon-gained-per-nitrogen-invested context.This study is dedicated to Professor M. Evenari for his continuing strong contributions to desert ecology     

青藏高原东部高山植物的水分利用效率与氮素利用效率研究

对生长在青藏高原东部隶属于23科、49属的71种高山植物(包括多年生和一年生植物)的稳定碳同位素比值、氮含量以及碳/氮比率进行了分析,并以稳定碳同位素比值及碳/氮比率来分别指示植物的水分利用效率和氮素利用效率.结果表明:(1)多年生植物稳定碳同位素比值显著高于一生年植物,而碳/氮比率显著低于一年生植物(P<0.01),氮含量两者无显著性差异.(2)多年生植物和一年生植物的稳定碳同位素比值均与碳/氮比率呈显著负相关(-0.643**和-0.707),而与氮含量均无明显相关性.研究证实,在自然条件下多年生植物的水分利用效率比一年生植物更高,而氮素利用效率却更低;高山植物水分利用效率和氮素利用效率存在明显的权衡",即植物不能同时提高水分利用效率和氮素利用效率,高水分利用效率的代价是降低氮素利用效率,青藏高原不同植物即使在相同环境条件下具有不同适应对策.     

Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

钠盐胁迫对藜科一年生草本植物生长和生物量分配的影响

选取河西走廊荒漠绿洲过渡带典型藜科一年生草本植物雾冰藜、刺沙蓬和白茎盐生草为研究对象,分析不同浓度盐分（NaCl和NaHCO₃,0、50、100、150、200 mmol/L）对3种藜科植物生长、繁殖和生物量分配的影响。研究结果表明：（1）钠盐胁迫下,3种藜科植物的存活率随盐浓度的增加呈下降趋势,雾冰藜和刺沙蓬在200 mmol/L NaCl和200 mmol/L NaHCO₃胁迫下无法存活或存活率极低,白茎盐生草在200 mmol/L NaHCO₃胁迫下无法存活;（2）钠盐胁迫显著抑制了刺沙蓬的生长和生物量积累,而一定浓度的盐分（50、100 mmol/L）可以促进雾冰藜和白茎盐生草的生长,较高浓度的盐分则抑制其生长;（3）3种植物的根冠比在钠盐胁迫下呈下降趋势,地上部生物量分配随盐浓度增加呈上升趋势,其中低盐胁迫下（50、100 mmol/L）繁殖分配比例增加明显,中高度盐胁迫下（150、200 mmol/L）茎、叶生物量分配比例增加显著,但根系生物量分配随盐分浓度增加而下降,这说明盐分胁迫下增加生物量在地上部的分配是藜科一年生草本植物应对盐胁迫的方式之一;（4） NaHCO₃的胁迫作用大于NaCl,3种植物中,白茎盐生草的耐盐性最强,而雾冰藜和刺沙蓬的耐盐能力较差。     

More plant biomass results in more offspring production in annuals,or does it?

Competitive ability in plants has been previously measured almost exclusively in terms of traits related to growth (biomass) or plant size. In this study, however, we used a multi‐species competition experiment with six annuals to measure relative competitive ability in terms of reproductive output, i.e. the number of offspring produced for the next generation. Under greenhouse conditions, plants of each species were started in pots from germinating seeds and were grown singly (free of competition) and at high density in both monocultures and in mixtures with all study species. Several traits traditionally regarded as determinants of competitive ability in plants were recorded for each species grown singly, including: seed mass, germination time, early growth rate and potential plant size (biomass and height). Under competition, several traits were recorded as indicators of relative performance in both monocultures and mixtures, including: biomass of survivors, total number of survivors, number of reproductive survivors, and reproductive output (total seed production) of the survivors. As expected, species that grew to a larger biomass in isolation had higher seed production in isolation. However, none of the traditional plant growth/size‐related traits, measured either in isolation or under competition, could predict between species variation in reproductive output under competition in either monocultures or mixtures. In mixtures, 97% of this variation in reproductive output could be explained by between‐species variation in the number of reproductive survivors. The results indicate that traits measured on plants grown singly may be poor predictors of reproductive output under competition, and that species’ rank order of competitive ability in terms of the biomass of survivors may bear no relationship to their rank order in terms of the number of offspring produced by these survivors. This has important implications for the interpretation of mechanisms of species coexistence and community assembly within vegetation.