Counteractive biomass allocation responses to drought and damage in the perennial herb Convolvulus demissus

Herbivory and water shortage are key ecological factors affecting plant performance. While plant compensatory responses to herbivory include reallocation of biomass from below‐ground to above‐ground structures, plant responses to reduced soil moisture involve increased biomass allocation to roots and a reduction in the number and size of leaves. In a greenhouse study we evaluated the effects of experimental drought and leaf damage on biomass allocation in Convolvulus demissus (Convolvulaceae), a perennial herb distributed in central Chile, where it experiences summer drought typical of Mediterranean ecosystems and defoliation by leaf beetles and livestock. The number of leaves and internode length were unaffected by the experimental treatments. The rest of plant traits showed interaction of effects. We detected that drought counteracted some plant responses to damage. Thus, only in the control watering environment was it observed that damaged plants produced more stems, even after correcting for main stem length (index of architecture). In the cases of shoot : root ratio, relative shoot biomass and relative root biomass we found that the damage treatment counteracted plant responses to drought. Thus, while undamaged plants under water shortage showed a significant increase in root relative biomass and a significant reduction in both shoot : root ratio and relative shoot biomass, none of these responses to drought was observed in damaged plants. Total plant biomass increased in response to simulated herbivory, apparently due to greater shoot size, and in response to drought, presumably due to greater root size. However, damaged plants under experimental drought had the same total biomass as control plants. Overall, our results showed counteractive biomass allocation responses to drought and damage in C. demissus. Further research must address the fitness consequences under field conditions of the patterns found. This would be of particular importance because both current and expected climatic trends for central Chile indicate increased aridity.     

Competitive reversals and environment-dependent resource partitioning in Erodium

Summary Effects of drought and varying plant density on the competitive coexistence of two winter annual Erodium species were studied using multiple regression analysis. Significant indications of resource partitioning were detected for interspecific mixtures under spring drought. Competitive superiority also was environment-dependent with E. botrys dominating with drought in autumn, while E. brachycarpum dominated with drought in spring. The results suggest that competitive coexistence in Erodium is promoted by processes both equilibrial (e.g., resource partitioning) and nonequilibrial (e.g. competitive reversals).     

Pre-mission invasion of Erodium cicutarium in California

Abstract. The California grassland is dominated by alien plant species. It is generally assumed that the invasion of aliens began with the initial introduction of livestock by Spanish missionaries in 1769. In this paper we present pollen evidence which indicates that Erodium cicutarium, a Mediterranean annual, was well established in the Santa Barbara region several years before the founding of the first California mission at San Diego in 1769. Historical evidence shows that it took the Spanish nearly a decade to develop a livestock base in California. Disturbance by livestock was therefore not a necessary prerequisite for invasion by alien plants. Historical and macrofossil evidence indicates that E. cicutarium invaded California from Baja California.     

PRODUCTION AND EFFECT OF SEED SIZE IN A GRASSLAND ANNUAL (ERODIUM BRACHYCARPUM,GERANIACEAE)

Over its fruit maturation period in the field, Erodium brachycarpum (Geraniaceae) exhibited a striking decrease in average seed mass; seeds produced in the 1st week of fruiting were twice the size of those produced 6 weeks later at the time of drought-induced plant senescence. Individuals with earliest fruit maturation produced the most seeds with the widest range of sizes and over a longer period than the others. In a greenhouse experiment, a similar decline in mean seed weight was generated by inducing drought conditions. Large seeds remained dormant longer and, once the seed coat was scarified, took longer to germinate than small seeds. This study indicates that small E. brachycarpum seeds (36% of the seed pool) produced late in the fruiting period are likely to germinate in the year of seed production, approximately 4–8 months after seed maturation. In contrast, the large seeds (23% of the seed pool), most of which are produced early in the fruiting period, are more likely to germinate in subsequent years (i.e., 15 months or more after seed maturation).     

Predicting the outcome of competition along a nitrogen gradient – a case study with Erodium cicutarium and Geranium pusillum

In order to understand the effect of a nitrogen gradient on the intraspecific and interspecific competition between plants, the two species Erodium cicutarium and Geranium pusillum were grown in a response-surface competion experiment at three different densities along a nitrogen gradient consisting of four different nitrogen levels. Seed set data were estimated from biomass measurements and fitted to a generalized hyperbolic competition model, and the probabilities of different ecological scenarios were calculated. The model predicted that E. cicutarium is competitively superior along most of the nitrogen gradient. This prediction was contrary to our prior expectations which were based on the occurance of the two species in natural habitats.     

Differential effect of drought regimes on the seedling performance of six floodplain grassland species

• The performance of seedlings is crucial for the survival and persistence of plant populations. Although drought frequently occurs in floodplains and can cause seedling mortality, studies on the effects of drought on seedlings of floodplain grasslands are scarce. We tested the hypotheses that drought reduces aboveground biomass, total biomass, plant height, number of leaves, leaf area and specific leaf area (SLA), and increases root biomass and root‐mass fraction (RMF) and that seedlings from species of wet floodplain grasslands are more affected by drought than species of dry grasslands.
• In a greenhouse study, we exposed seedlings of three confamilial pairs of species (Pimpinella saxifraga, Selinum carvifolia, Veronica teucrium, Veronica maritima, Sanguisorba minor, Sanguisorba officinalis) to increasing drought treatments. Within each plant family, one species is characteristic of wet and one of dry floodplain grasslands, confamilial in order to avoid phylogenetic bias of the results.
• In accordance with our hypotheses, drought conditions reduced aboveground biomass, total biomass, plant height, number of leaves and leaf area. Contrary to our hypotheses, drought conditions increased SLA and decreased root biomass and RMF of seedlings. Beyond the effects of the families, the results were species‐specific (V. maritima being the most sensitive species) and habitat‐specific. Species indicative of wet floodplain grasslands appear to be more sensitive to drought than species indicative of dry grasslands.
• Because of species‐ and habitat‐specific responses to reduced water availability, future drought periods due to climate change may severely affect some species from dry and wet habitats, while others may be unaffected.

     

长江口九段沙湿地海三棱藨草生物量分配特征及其影响因子

海三棱藨草(Scirpus mariqueter)为长江河口盐沼湿地先锋植物。以长江口九段沙湿地为主要研究区域,在江亚南沙、上沙、中下沙等不同区域设置固定站点,进行植被和环境因子的取样调研,分析研究了海三棱藨草的生物量分配特征及其主要影响因子,以期为海三棱藨草的培育与恢复、盐沼湿地生态系统的保护与利用提供科学依据。结果表明:(1)不同站点海三棱藨草的茎基高度、球茎生物量占比、根茎与球茎之比均不存在显著差异(P0.05),而植株密度、植株高度、根茎生物量占比、果实生物量占比、茎叶生物量占比、须根生物量占比、果实与根茎之比、果实与球茎之比在不同站点间存在显著差异(P0.05);(2)不同站点的沉积物电导率、盐度、容积密度(5—10、25—30 cm土层)、总碳含量(10—15、20—25、25—30 cm土层)、总氮含量(5—10 cm土层)存在显著差异(P0.05);(3)海三棱藨草的果实与根茎的生物量之比和沉积物5—10、25—30 cm总碳含量,0—5、5—10 cm总氮含量,0—5 cm容积密度,茎基高度6种因子的组合存在极显著相关(P0.01);而其他生物量分配指标与本文涉及到的环境因子、植物表形参数均不存在显著相关性(P0.05)。海三棱藨草生物量分配特征是其自身生长特征与环境因子综合作用的结果。在今后的研究中需拓展生境因子涵盖范围,深入分析多因子综合作用对海三棱藨草生物量分配特征的影响。     

Phenolics Composition and Biological Activities Assessment of Leaves,Flowers and Roots Extracts from Erodium glaucophyllum,Erodium hirtum and Erodium guttatum

Biology Bulletin - Erodium glaucophyllum, Erodium hirtum and Erodium guttatum were medicinal herbs from a southern Mediterranean known for its health benefits. There is a strong demand for the find...     

Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7 years (1999–2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area.     

Recent loss of plastid-encoded ndh genes within Erodium (Geraniaceae)

Plastid genomes in the flowering plant family Geraniaceae are known to be highly rearranged based on complete sequences representing the four major genera Erodium, Geranium, Monsonia, and Pelargonium. In this paper we report on the genome sequence of a second species of Erodium, E. carvifolium, representing the second major clade (clade II) in the phylogeny of this genus. Comparison of this genome sequence to the previously published sequence of E. texanum from clade I demonstrates that the plastid genomes of these two species encode the same number of proteins but differ greatly in their relative degree of rearrangement; 14 kb of additional sequence in E. texanum contains complex repeats associated with rearrangement endpoints, whereas the plastid genome of E. carvifolium is streamlined at 116 kb and displays no unique alterations in gene order. Furthermore, these species from both major clades of Erodium contain intact NADH dehydrogenase (ndh) genes, but the 11 ndh genes are represented as pseudogenes in a small clade of 13 species. It is unclear whether plastid-encoded ndh genes have been lost entirely or functionally transferred to the nucleus. This is the third report of the absence of functional ndh genes, and the current study describes the most recent loss of these genes among photosynthetic seed plants and the second such loss among angiosperms. The other ndh losses from Pinaceae/Gnetales and Orchidaceae are much more ancient. Comparative biochemistry between Erodium species with and without plastid-encoded ndh genes may elucidate changes in photosynthetic function and the role of the Ndh complex.     

PARTITIONING OF SPACE AND WATER BETWEEN TWO CALIFORNIA ANNUAL GRASSLAND SPECIES

We examined the potential for resource partitioning between two sympatric species with similar phenologies but different rooting morphologies. The annual grass, Bromus diandrus (Roth.), and annual forb, Erodium botrys (Cav.), were grown in monoculture and 50:50 mixed stands at each of three densities (10, 30, 100 seeds/dm²) in a randomized complete block design. Plants were grown outdoors, in 1-m-tall 15-cm-diameter containers. Comparison of seed number produced per plant in mixture and in monoculture indicated greater effects of intraspecific than interspecific competition for Erodium. Such differences were not detected for Bromus seed number, and the converse relationship was suggested from Bromus shoot biomass. Final size inequality of Bromus populations tended to be higher in monoculture than in mixture; no patterns in Erodium size distribution over time or stand composition were evident. Bromus roots were primarily in the upper 10 cm of soil, while Erodium roots were bimodally distributed in the surface and deep soil. Roots of the two species in mixture showed a distribution pattern intermediate between those of the two monocultures. The rate of soil water depletion was higher in the high density than in the low density stands, but was not dependent on stand composition at a given density. Partitioning of belowground space and water resources by groups of species with different root morphologies may partially explain the high species diversity in the grassland.     

Interactive effects of elevated temperature and drought on plant carbon metabolism: A meta-analysis

Elevated temperature (T_e) and drought often co-occur and interactively affect plant carbon (C) metabolism and thus the ecosystem C cycling; however, the magnitude of their interaction is unclear, making the projection of global change impacts challenging. Here, we compiled 107 journal articles in which temperature and water availability were jointly manipulated, and we performed a meta-analysis of interactive effects of T_e and drought on leaf photosynthesis (A_growth) and respiration (R_growth) at growth temperature, nonstructural carbohydrates and biomass of plants, and their dependencies on experimental and biological moderators (e.g., treatment intensity, plant functional type). Our results showed that, overall, there was no significant interaction of T_e and drought on A_growth. T_e accelerated R_growth under well-watered conditions rather than under drought conditions. The T_e × drought interaction on leaf soluble sugar and starch concentrations were neutral and negative, respectively. The effect of T_e and drought on plant biomass displayed a negative interaction, with T_e deteriorating the drought impacts. Drought induced an increase in root to shoot ratio at ambient temperature but not at T_e. The magnitudes of T_e and drought negatively modulated the T_e × drought interactions on A_growth. Root biomass of woody plants was more vulnerable to drought than that of herbaceous plants at ambient temperature, but this difference diminished at T_e. Perennial herbs exhibited a stronger amplifying effect of T_e on plant biomass in response to drought than did annual herbs. T_e exacerbated the responses of A_growth and stomatal conductance to drought for evergreen broadleaf trees rather than for deciduous broadleaf and evergreen coniferous trees. A negative T_e × drought interaction on plant biomass was observed on species-level rather than on community-level. Collectively, our findings provide a mechanistic understanding of the interactive effects of T_e and drought on plant C metabolism, which would improve the prediction of climate change impacts.     

Increased Reproductive Allocation of Rumex japonicus Caused by Leaf Clipping

Abstract To elucidate the effects of herbivory by chrysomelid beetles on Rumex japonicus, rosette leaves were clipped and the subsequent fruit production and root growth were observed. The increase of leaf biomass of some clipped plants was greater than that of control plants, although this varied among individual plants. The root growth of clipped plants was less than that of control plants. Fruit production increased with plant size, and there was no difference in fruit production between clipped and control plants. Reproductive allocation (fruit biomass, relative to fruit biomass plus root growth) increased with plant size; it was greater in clipped plants than in control ones. Based on these results, reproductive allocation strategy against herbivory was discussed.     

干旱胁迫对欧李幼苗表型可塑性的影响

该研究以欧李为材料,探讨了干旱胁迫对欧李表型可塑性的影响。结果表明:(1)随着干旱胁迫的加剧,欧李根生物量、枝叶生物量、植株总生物量积累、根冠比和根冠比胁迫指数均呈现先升高后降低的趋势,在T1处理下达到最大值,并显著高于其他处理(α=0.05)。(2)随土壤含水量的降低,欧李根的生物量分配指数呈现先增加后降低的趋势,叶生物量与之相反,在T1处理下根的生物量分配指数最大,枝叶的最小(α=0.05)。(3)在水分供应为60%～80%时,欧李的株高、冠幅、基径、二级分枝数、主根长、主根直径及侧根数量均达到最大值(α=0.05),对一级分枝数的生长没有显著影响。(4)随着水分胁迫的加剧,叶片长从T2处理开始下降,叶片宽、单片叶面积及比叶面积均呈现先增加后减少的趋势(α=0.05)。综上可得,欧李通过调整形态特性和各器官生物量积累及其分配对不同干旱胁迫条件产生了较强的可塑性。     

Drought affects the coordination of belowground and aboveground resource‐related traits in Solidago canadensis in China

Quantifying patterns of variation and coordination of plant functional traits can help to understand the mechanisms underlying both invasiveness and adaptation of plants. Little is known about the coordinated variations of performance and functional traits of different organs in invasive plants, especially in response to their adaptation to environmental stressors. To identify the responses of the invasive species Solidago canadensis to drought, 180 individuals were randomly collected from 15 populations and 212 ramets were replanted in a greenhouse to investigate both the response and coordination between root and leaf functional traits. Drought significantly decreased plant growth and most of the root and leaf functional traits, that is, root length, surface area, volume and leaf size, number, and mass fraction, except for the root length ratio and root mass fraction. Phenotypic plasticity was higher in root traits than in leaf traits in response to drought, and populations did not differ significantly. The plasticity of most root functional traits, that is, root length (RL), root surface area (RSA), root volume (RV), and root mass fraction (RMF), were significantly positively correlated with biomass between control and drought. However, the opposite was found for leaf functional traits, that is, specific leaf area (SLA), leaf area ratio (LAR), and leaf mass fraction (LMF). Drought enhanced the relationship between root and leaf, that is, 26 pairwise root–leaf traits were significantly correlated under drought, while only 15 pairwise root–leaf traits were significantly correlated under control conditions. Significant correlations were found between biomass and all measured functional traits except for leaf size. RV, root length ratio, RMF, total area of leaves, and LMF responded differently to water availability. These responses enable S. canadensis to cope with drought conditions and may help to explain the reason of the vast ecological amplitude of this species.     

Patterns of resource allocation in the dioecious alpine herb Aciphylla simplicifolia (Apiaceae)

Abstract Patterns of reproductive and vegetative biomass allocation were compared in male and female plants of the alpine herb Aciphylla simplicifolia. Male and female plants had similar vegetative biomass but differed in the pattern of resource allocation. Inflorescences of males and females were similar in weight at the time of flowering, but differed in biomass allocation to some structures within the inflorescences, particularly those associated with ovule production and pollinator attraction (number and size of flowers). At the time of fruit production, female inflorescences were 2.6 times heavier than at flowering with developing fruit six times heavier than flowers. In addition to the increase in biomass allocated to structures associated with the provisioning and dissemination of seed, support structures (main and side stalks) were also heavier. As a result of this additional investment of resources at the time of fruit production, the reproductive effort (RE) of female plants was much higher than that of males: 37% of above ground biomass compared with 21% for males. Differences in RE did not change with plant size; however, allocation to reproduction appeared to be a constant proportion of biomass over nearly all plant sizes sampled. These results show that sex‐specific resource allocation can be a complex of temporal and morphological patterns.     

Plant size influences abundance of floral visitors and biomass allocation for the cushion plant Thylacospermum caespitosum under an extreme alpine environment

Variation in size may influence the abundance of visitors and reproductive allocation for cushion plants in the extreme alpine environments. To assess effects of plant size on the abundance of main visitors and reproductive allocation in Thylacospermum caespitosum populations at two altitudes, the abundance of the visitors, visiting frequency, total number of flowers, number of fruits, number of unseeded flowers, and reproductive allocation were investigated during the period of reproductive growth. Concurrently, the effects of plant size on the visitors' contributions to fruit setting rate were assessed by a bagging experiment. Our results showed that flies (Musca domestica and Dasyphora asiatica) were the main pollinating insects of T. caespitosum, and they could obvious facilitate (p < 0.05) the fruit setting rate of this cushion plant. Seed set and floral visitation were significantly influenced (p < 0.001) by plant size. Moreover, the reproductive allocation and fruit setting rate of T. caespitosum was influenced (p < 0.001) by plant size. More biomass was allocated to reproduction in plants of greater diameter. There is an increase in reproductive success (increases of fruit number with increase in plant size) in relation to plant size. In conclusion, the extent of M. domestica and D. asiatica to facilitate the fruit setting rate mainly depended on the size of T. caespitosum. Size‐dependent reproductive allocation occurred in T. caespitosum and was the chief factor affecting the contribution of flies to fruit setting rate. These traits reflect reproductive fitness of T. caespitosum related to plant size in extreme alpine environments.     

Effects of the Lycopersicon chmielewskii sucrose accumulator gene (sucr) on fruit yield and quality parameters following introgression into tomato

A gene controlling fruit sucrose accumulation, sucr, was introgressed from the wild tomato species Lycopersicon chmielewskii into the genetic background of a hexose-accumulating cultivated tomato, L. esculentum. During introgression, the size of the L. chmielewskii chromosomal segment containing sucr was reduced by selection for recombination between RFLP markers for the sucr gene and flanking loci. The effects of sucr on soluble solids content, fruit size, yield and other fruit parameters were studied in the genetic background of the processing tomato cultivar Huntl00. In a segregating BC₅F₂ generation, the smallest introgression containing sucr-associated markers was necessary and sufficient to confer high-level sucrose accumulation, the effects of which were completely recessive. Fruit of sucr/sucr genotypes were smaller than those of +/sucr or +/+ genotypes at all stages of development. The timing of sugar accumulation and total sugar concentration were unaffected by sugar composition. No differences in total fruit biomass (fresh weight of red and green fruit) at harvest were observed between the genotypes, and sucrose accumulators produced greater numbers of fruit than hexose accumulators in one family. However, the proportion of ripe fruit at harvest, and hence yield of ripe fruit, as well as average ripe fruit weight and seed set were reduced in sucr/sucr genotypes. Sucrose accumulation was also associated with increased soluble solids content, consistency, serum viscosity, predicted paste yield and acidity, and decreased color rating. In the first backcross to L. chmielewskii, hexose accumulators (+/sucr) had larger fruit than sucrose accumulators (sucr/sucr), while no difference in soluble solids was detected.     

Protozoa,Nematoda and Lumbricidae in the rhizosphere of Hordelymus europeaus (Poaceae): faunal interactions,response of microorganisms and effects on plant growth

Interactions among protozoa (mixed cultures of ciliates, flagellates and naked amoebae), bacteria-feeding nematodes (Pellioditis pellio Schneider) and the endogeic earthworm species Aporrectodea caliginosa (Savigny) were investigated in experimental chambers with soil from a beechwood (Fagus sylvatica L.) on limestone. Experimental chambers were planted with the grass Hordelymus europeaus L. (Poaceae) and three compartments separated by 45-m mesh were established: rhizosphere, intermediate and non-rhizosphere. The experiment lasted for 16 weeks and the following parameters were measured at the end of the experiment: shoot and root mass of H. europaeus, carbon and nitrogen content in shoots and roots, density of ciliates, amoebae, flagellates and nematodes, microbial biomass (SIR), basal respiration, streptomycin sensitive respiration, ammonium and nitrate contents, phosphate content of soil compartments. In addition, leaching of nutrients (nitrogen and phosphorus) and leachate pH were measured at regular intervals in leachate obtained from suction cups in the experimental chambers. Protozoa stimulated the recovery of nitrifying bacteria following defaunation (by chloroform fumigation) and increased nitrogen losses as nitrate in leachate. In contrast, protozoa and nematodes reduced leaching of phosphate, an effect ascribed to stimulation of microbial growth early in the experiment. Earthworms strongly increased the amount of extractable mineral nitrogen whereas it was strongly reduced by protozoa and nematodes. Both protozoa and nematodes reduced the stimulatory effect of earthworms on nitrogen mineralization. Microbial biomass, basal respiration, and numbers of protozoa and nematodes increased in the vicinity of the root. Protozoa generally caused a decrease in microbial biomass whereas nematodes and earthworms reduced microbial biomass only in the absence of protozoa. None of the animals studied significantly affected basal respiration, but specific respiration of microorganisms (O₂ consumption per unit biomass) was generally higher in animal treatments. The stimulatory effect of nematodes and earthworms, however, occurred only in the absence of protozoa. The sensitivity of respiration to streptomycin suggested that protozoa selectively grazed on bacterial biomass but the bacterial/fungal ratio appeared to be unaffected by grazing of P. pellio. Earthworms reduced root biomass of H. europaeus, although shoot biomass remained unaffected, and concentrations of nitrogen in shoots and particularly in roots were strongly increased by earthworms. Both nematodes and protozoa increased plant biomass, particularly that of roots. This increase in plant biomass was accompanied by a marked decrease in nitrogen concentrations in roots and to a lesser extent in shoots. Generally, the effects of protozoa on plant growth considerably exceeded those of nematodes. It is concluded that nematodes and protozoa stimulated plant growth by non-nutritional effects, whereas the effects of earthworms were caused by an increase in nutrient supply to H. europaeus.     

Genome‐wide association of drought‐related and biomass traits with HapMap SNPs in Medicago truncatula

Improving drought tolerance of crop plants is a major goal of plant breeders. In this study, we characterized biomass and drought‐related traits of 220 Medicago truncatula HapMap accessions. Characterized traits included shoot biomass, maximum leaf size, specific leaf weight, stomatal density, trichome density and shoot carbon‐13 isotope discrimination (δ¹³C) of well‐watered M. truncatula plants, and leaf performance in vitro under dehydration stress. Genome‐wide association analyses were carried out using the general linear model (GLM), the standard mixed linear model (MLM) and compressed MLM (CMLM) in TASSEL, which revealed significant overestimation of P‐values by CMLM. For each trait, candidate genes and chromosome regions containing SNP markers were found that are in significant association with the trait. For plant biomass, a 0.5 Mbp region on chromosome 2 harbouring a plasma membrane intrinsic protein, PIP2, was discovered that could potentially be targeted to increase dry matter yield. A protein disulfide isomerase‐like protein was found to be tightly associated with both shoot biomass and leaf size. A glutamate‐cysteine ligase and an aldehyde dehydrogenase family protein with Arabidopsis homologs strongly expressed in the guard cells were two of the top genes identified by stomata density genome‐wide association studies analysis.