Year-to-year changes in expression of maternal effects in perennial plants

Although there are numerous examples of maternal effects in perennial plant species, studies usually follow the fate of progeny only in their juvenile stages or for one growing season. Here we experimentally demonstrate, for two perennial species, that maternal nutrient environments and disturbance histories affect progeny differently during their first two growing seasons.Whereas progeny of mothers that suffered nutrient insufficiency produced more spikes in the first season, they produced equal spikes in the second season when compared to progeny of mothers from benign conditions. The progeny of mothers that had been grown in nutrient-poor conditions grew longer leaves in their second year, when compared to progeny of mothers from nutrient-poor conditions that experienced severe disturbance (removal of all above-ground biomass) but this was not the case in their first year. Additionally, progeny of mothers that experienced severe disturbance and were grown in nutrient-poor conditions produced longer leaves when compared to progeny of disturbed mothers grown in nutrient-rich conditions in the second year but this pattern was not observed in first year of the study.The changing expression of maternal effects in our study showed the necessity of longer-term studies to identify the effects and to determine their roles in the ecology of perennial species. We also suggest possible mechanisms responsible for the observed patterns.     

Maternal effects alter progeny's response to disturbance and nutrients in two Plantago species

Environmental stress leading to a decrease in growth may be compensated for later in ontogeny by a growth plastic response. Such response could be also transmitted to the next generation, which is called transgenerational plasticity. In this study, two Plantago species were used to test whether compensation for biomass loss after disturbance is driven by maternal effects (ME) due to nutrients and disturbance, i.e. by the form of transgenerational plasticity. Additionally, we tested whether ME could contribute to a different performance of progeny having different disturbance histories. We also tested whether ME are adaptive and whether they differ between related species. The degree of (over)compensation for biomass loss was affected by ME. Maternal effects resulted in different performance of disturbed over undisturbed progeny in relation to nutrient status of the progeny environment along with disturbance and nutrients experienced by mothers. Progeny of P. lanceolata grew more leaf biomass when grown in the same nutrient conditions as experienced by their mothers suggesting that maternal effects might be adaptive. Although overall, there was a consistent role of ME in biomass compensation after disturbance among the two species, there were also some species‐specific effects. We conclude that compensation for biomass loss is driven both by maternal effects and by progeny environment. This may lead to the different success of regenerative strategies in environments with contrasting nutrient levels. The different role of ME even between related species may contribute to ecological diversity among species.     

Effect of support availability,mother plant genotype and maternal support environment on the twining vine <Emphasis Type="Italic">Ipomoea purpurea</Emphasis>

In the twining vine Ipomoea purpurea we experimentally assessed the effect of support availability (a vertical stake), maternal genotype (family), and maternal environment (presence or absence of support in mother plants) on morphological traits, accounting for differences in initial seed size. While there was no effect of the maternal environment on seed size at the family level, families within each maternal environment did differ in seed size. Seeds from families of supported mothers showed higher germination than those from families of unsupported mothers. The maternal environment did not influence shoot traits but affected phenotypic plasticity in the number of leaves in response to support. Thus, whereas progeny plants from unsupported mothers did not show a response to support availability in the number of leaves, progeny from supported mother plants had a greater number of leaves once support was provided. The maternal genotype only affected the number of leaves.     

Adaptive transgenerational plasticity in the perennial Plantago lanceolata

Phenotypes of plants, and thus their ecology and evolution, can be affected by the environmental conditions experienced by their parents, a phenomenon called parental effects or transgenerational plasticity. However, whether such effects are just passive responses or represent a special type of adaptive plasticity remains controversial because of a lack of solid tests of their adaptive significance. Here, we investigated transgenerational effects of different nutrient environments on the productivity, carbon storage and flowering phenology of the perennial plant Plantago lanceolata, and whether these effects are influenced by seasonal variation in the maternal environment. We found that maternal environments significantly affected the offspring phenotype, and that plants consistently produced more biomass and had greater root carbohydrate storage if grown under the same environmental conditions as experienced by their mothers. The observed transgenerational effects were independent of the season in which seeds had matured. We therefore conclude that transgenerational effects on biomass and carbon storage in P. lanceolata are adaptive regardless of the season of seed maturation.     

Maternal and grandmaternal age influence offspring fitness in Drosophila.

The influence of maternal and grandmaternal age on progeny egg-to-adult viability was assessed in Drosophila serrata. Viability in progeny decreased with increasing maternal age. The potential for cumulative age effects was investigated in two environments, one of which involved nutrient and cold stress. Environment influenced viability and female age influenced progeny egg-to-adult viability across one generation. The influence on viability was cumulative across two generations. Females from old mothers, who also had old grandmothers, had the lowest viability in both environments. Grandmaternal effects were associated with a decrease in egg hatch rate whereas maternal effects also involved larval-to-adult viability. The age of the mother and grandmother should be taken into account when evaluating life-history traits in Drosophila.     

EFFECT OF THE PARENTAL NUTRIENT REGIME ON GROWTH OF THE PROGENY IN ABUTILON THEOPHRASTI (MALVACEAE)

Two genets of Abutilon theophrasti were clonally replicated and grown to maturity in a glasshouse at two levels of nutrient supply. The seeds produced were weighed and the resulting seedlings exposed to two nutrient levels during development. The progeny plants were harvested on three occasions, and leaf areas and dry weights of the different plant parts were determined. At the early stages of growth an increase in the maternal nutrient supply significantly increased seedling height, cotyledon area, and leaf areas, and seed weight had a significant effect on several traits. The maternal nutrient addition had no significant effect at a later stage of growth (35 d after sowing), but at 56 d after sowing it did affect offspring leaf areas and dry weights. Significant interaction terms indicate that the response to parental nutrient addition may depend both on genotype and on the nutrient status of the progeny. Different plant characters are differentially sensitive to maternal conditions and these may be expressed at different stages of development.     

Systemic regulation of leaf anatomical structure, photosynthetic performance, and high-light tolerance in sorghum

Leaf anatomy of C3 plants is mainly regulated by a systemic irradiance signal. Since the anatomical features of C4 plants are different from that of C3 plants, we investigated whether the systemic irradiance signal regulates leaf anatomical structure and photosynthetic performance in sorghum (Sorghum bicolor), a C4 plant. Compared with growth under ambient conditions (A), no significant changes in anatomical structure were observed in newly developed leaves by shading young leaves alone (YS). Shading mature leaves (MS) or whole plants (S), on the other hand, caused shade-leaf anatomy in newly developed leaves. By contrast, chloroplast ultrastructure in developing leaves depended only on their local light conditions. Functionally, shading young leaves alone had little effect on their net photosynthetic capacity and stomatal conductance, but shading mature leaves or whole plants significantly decreased these two parameters in newly developed leaves. Specifically, the net photosynthetic rate in newly developed leaves exhibited a positive linear correlation with that of mature leaves, as did stomatal conductance. In MS and S treatments, newly developed leaves exhibited severe photoinhibition under high light. By contrast, newly developed leaves in A and YS treatments were more resistant to high light relative to those in MS- and S-treated seedlings. We suggest that (1) leaf anatomical structure, photosynthetic capacity, and high-light tolerance in newly developed sorghum leaves were regulated by a systemic irradiance signal from mature leaves; and (2) chloroplast ultrastructure only weakly influenced the development of photosynthetic capacity and high-light tolerance. The potential significance of the regulation by a systemic irradiance signal is discussed.     

The Detection,Rate and Manifestation of Residual Sexuality in Apomictic Populations of Pilosella (Asteraceae,Lactuceae)

The effect of maternal, facultatively apomictic plants on population diversity was evaluated in seven hybridizing polyploid Pilosella populations, where apomictic (P. bauhini or P. aurantiaca) and sexual (P. officinarum) biotypes coexist. The ploidy level, reproductive system, morphology, clonal structure and chloroplast DNA haplotypes were used to characterize these plants and their hybrids. The reproductive origins of the progeny were assessed through either a flow cytometric seed screen and/or a comparison between the ploidy level of progeny embryos/seedlings and the maternal ploidy level. The cultivated progeny derived from residual sexuality in maternal apomicts were also identified based on their morphology and reproductive behaviour. The progeny different from their maternal parents (0.6?92.3 % of progeny embryos and 0?100 % of progeny seedlings) originated either sexually or via haploid parthenogenesis. Comparing the facultatively apomictic and sexual mothers, the progeny arrays generated in the field showed that apomictic mothers produce progeny that is more variable in ploidy level. This effect was demonstrated at both the embryonic and seedling stages of progeny development. Residual sexuality in apomicts was also effective in experimental crosses, generating progeny similar to spontaneous hybrids in the field. The 2n + n hybrids produced from an apomictic and a sexual parent displayed similar reproductive behaviour, producing polyhaploid, sexual and apomictic progeny in variable ratios. Repeated hybridizations between parental species and/or multi-step crosses can result in hybrid swarms rich in cytotypes and morphotypes. The variation recorded in these populations suggests prevailing introgressive hybridization towards the sexual species P. officinarum.     

Enriching Ploidy Level Diversity: the Role of Apomictic and Sexual Biotypes of Hieracium subgen. Pilosella (Asteraceae) that Coexist in Polyploid Populations

The capacity to generate variation in ploidy and reproductive mode was compared in facultatively apomictic versus sexual maternal plants that coexist in two model populations. The population structure was studied in polyploid hybrid swarms comprised of Hieracium pilosella (usually sexual, less commonly apomictic), H. bauhini (apomictic), and their hybrids (sexual, apomictic, or sterile). Relationships among established biotypes were proposed on the basis of their DNA ploidy level/chromosome number, reproductive mode and morphology. Isozyme phenotypes and chloroplast DNA haplotypes were assayed in the population that was richer in hybrids. The reproductive origin of seed progeny was identified in both sexual and apomictic mothers, using alternative methods: the karyological, morphological and reproductive characters of the cultivated progeny were compared with those of respective mothers, or flow cytometric seed screening was used. In both populations, the progeny of sexual mothers mainly retained a rather narrow range of ploidy level/chromosome number, while the progeny of facultatively apomictic mothers was more variable. The high-polyploid hybrids, which had arisen from the fertilization of unreduced egg cells of apomicts, mainly produced aberrant non-maternal progeny (either sexually and/or via haploid parthenogenesis). Apparently, such versatile reproduction resulted in genomic instability of the recently formed high-polyploid hybrids. While the progeny produced by both true apomictic and sexual mothers mostly maintained the maternal reproductive mode, the progeny of those ‘versatile’ mothers was mainly sexual. Herein, we argue that polyploid facultative apomicts can considerably increase population diversity.     

Maternal effects of drought stress and inbreeding in Impatiens capensis (Balsaminaceae)

Maternal effects can have substantial impacts on plant fitness and plant populations. Stressful environmental conditions can cause a maternal plant to inadequately provision its progeny, resulting in poor seedling growth, low reproductive success, and decreased competitive ability. Maternal effects consist of environmental and genetic load components, but the interactions between these two components have rarely been considered. To determine the effects of maternal drought stress and maternal inbreeding on progeny biomass (a fitness correlate) and physiological responses to drought stress, we conducted a greenhouse experiment with genetic lines from two populations (mesic site vs. dry site) of the herbaceous annual Impatiens capensis (Balsaminaceae). Seeds were collected from cleistogamous flowers of inbred or outcrossed maternal plants that were subject to either a drought or control treatment. These seeds were grown into juvenile plants that were also subject to either a drought stress or a control treatment. Plants from the mesic site had significantly reduced biomass from maternal drought stress, while plants from the dry site maintained biomass despite adverse maternal environmental conditions. Juvenile plants of both populations had reduced biomass only as a result of maternal inbreeding. Interestingly, inbreeding depression was more apparent when maternal environmental conditions were benign.     

硅和干旱胁迫对水稻叶片光合特性和矿质养分吸收的影响

硅被认为是植物生长的有益元素,它能增强植物对非生物逆境和生物逆境胁迫的抗性。以抗旱性不同的一对水稻近等基因系w-14-和w-20为实验材料,采用盆栽实验,研究了干旱胁迫下硅处理对水稻生长性状、光合生理特性和矿质养分吸收的影响。结果表明,在正常水分条件下硅处理对水稻的生长及生理特性没有明显影响。干旱胁迫显著降低水稻植株的生长,叶绿素含量、叶绿素荧光参数Fv/Fm及Fv/F0值显著降低,光合作用受到明显抑制。加硅能提高干旱胁迫条件下水稻植株的生物量、水分利用效率、叶片叶绿素含量、净光合速率和蒸腾速率,而气孔导度和细胞间隙CO2浓度则下降。无论干旱与否,施硅后水稻的叶片硅含量均显著上升。两个水稻品系叶片的无机离子含量在干旱胁迫条件下均呈显著增加的趋势,而硅处理后材料w-14的叶片K+、Na+、Ca2+、Mg2+、Fe3+含量分别降低16.38%,24.50%,19.70%,21.52%,18.58%,w-20则分别降低11.64%,12.11%,16.06%,11.11%和19.15%,并使之回复到与对照更接近的水平。研究结果表明了硅提高水稻植株的抗旱性与光合作用的改善和矿质养分的调节有关。     

Impacts of eriophyoid gall mites on arctic willow in a rapidly changing Arctic

Arctic plants and herbivores are subject to ongoing climatic changes that are more rapid and extreme than elsewhere on the planet, and thus it is pivotal to understand the arctic plant-herbivore interactions in a global change context. We examined how infestation by an eriophyoid gall mite affects the circumpolar shrub Salix arctica, and how the effects vary across vegetation types. Specifically, we compared multiple leaf characteristics (leaf area, biomass, nutrient levels, δ¹⁵N and δ¹³C, and stress and performance of the photosynthetic apparatus) of infested leaves to those of un-infested leaves. Furthermore, we examined how altered environmental conditions, here experimentally manipulated levels of temperature, water and nutrients, shading, and UV-B radiation, affect the prevalence, density, and intensity of gall mite infestation and its impacts on S. arctica. Infested leaves were smaller in area and biomass and had lower nitrogen and carbon pools. However, their carbon concentration was higher, possibly because the galls acted as carbon sinks. The smaller photosynthetic area and lower nutrient content caused increased stress on the photosynthetic apparatus in infested leaves. The remaining leaf tissue responded with a higher photosynthetic performance, although there were indications of a general reduction in photosynthesis. Female leaves were more affected than male leaves. The experimental manipulations of environmental conditions did not affect the gall prevalence, density, or intensity on S. arctica leaves. Rather, plants responded positively to the treatments, reducing the effects of the galls to in-significance. This suggests a higher tolerance and defense against gall mites under future climate conditions.     

Rethinking the thermal melanism hypothesis: rearing temperature and coloration in pygmy grasshoppers

Selection for efficient conversion of solar radiation to body heat has favored the evolution of dark coloration in many ectotherms. The thermal melanism hypothesis posits that dark coloration is beneficial under conditions of low ambient temperatures because it results in faster heating rates and higher body temperatures. Fast heating rates, however, may come at a cost of overheating unless compensated for by thermal physiology or behaviour. Pygmy grasshopper (Orthoptera, Tetrigidae) populations that inhabit fire-ravaged areas characterized by blackened backgrounds and hot surface temperatures due to high absorbance of solar radiation show an increased frequency of black phenotypes. I raised the progeny of wild-captured Tetrix undulata in cold and hot temperatures and used data on color patterns and survival in a greenhouse to examine whether a cold thermal environment triggered the development of melanic coloration or differently affected survival of melanic versus non-melanic individuals. My results indicate that melanism was not influenced by rearing temperature but by genes or epigenetic maternal effects. Temperature also did not affect survival. However, melanic individuals produced by melanic mothers survived longer than melanic individuals produced by non- melanic mothers, whereas non-melanic individuals produced by non-black mothers survived longer than melanic individuals produced by non-black mothers. This suggests a mismatch between color and physiology in offspring belonging to a different color morph than their mother. Future investigations into the evolution of melanism should consider conflicting selection pressures on thermal capacity and camouflage as well as the influence of correlated responses to selection on traits associated with coloration.     

Effect of mycorrhizal-enhanced leaf phosphate status on carbon partitioning, translocation and photosynthesis in cucumber

An assessment of the effects of arbuscular mycorrhizal (AM) infection on photosynthesis, carbon (C) allocation, translocation and biomass production of cucumber, grown in sand culture, was made using a previously determined phosphorus (P) supply (0·13 mol m^?3 P) which had a significant impact on AM infection. Separation of a direct effect of AM infection from an indirect one due to an enhanced leaf P status was achieved using a comparable non‐mycorrhizal treatment (NAM + P) supplemented with extra P (0·19 mol m^?3 P). Total leaf P concentration, specific leaf mass, photosynthetic capacity, and incorporation of ¹⁴C into non‐structural carbohydrate pools were dependent on leaf age. Both maximum and ambient photosynthetic rates were significantly higher in the youngest fully expanded leaves from AM and NAM + P plants which also had the higher leaf P concentrations. There were no differences in the total concentrations of starch, sucrose, raffinose or stachyose in young or old leaves among AM, non‐mycorrhizal (NAM) and NAM + P treatments. However, younger leaves of NAM plants showed a shift in ¹⁴C‐partitioning from stachyose and raffinose synthesis to starch accumulation. Determination of ADP‐glucose pyrophosphorylase (AGPase), sucrose synthase and sucrose phosphate synthase enzyme activities revealed that only AGPase activity was correlated with the increased incorporation rate of ¹⁴C into starch in young leaves of NAM plants. Although there were significant AM‐specific effects on C translocation to the root system, AM plants had similar rate of photosynthesis to NAM + P plants. These results suggest that the increase in photosynthetic rate in leaves of AM‐infected cucumber was due to an increased P status, rather than a consequence of a mycorrhizal ‘sink’ for assimilates.     

Leaf gas exchange of cassava as affected by quality of planting material and water stress

Field trial was conducted to study the effects of quality of planting material and prolonged water stress on leaf gas exchange of the cassava (Manihot esculenta Crantz) cultivar M Col 1684. Nutrient contents of planting material affected rootlet formation, but not leaf gas exchange. Net photosynthetic rate (PN), stomatal conductance (gs), and intercellular CO2 concentration (Ci) were significantly reduced by prolonged water stress. New leaves developed after recovery from water stress showed higher PN and gs, as compared to leaves of similar ages of unstressed plants. The higher PN was associated with higher leaf nutrient contents, indicating that photosynthetic capacity was enhanced in these leaves. These compensating characteristics may partly explain the small yield reduction often observed in stressed cassava.     

Analyzing the significance of defoliation in growth, photosynthetic compensation and source-sink relations

Leaf canopy plays a determining role influencing source-sink relations as any change in source activity (photosynthesis) affects sink metabolism. Defoliation (removal of leaves) influences growth and photosynthetic capacity of plants, remobilizes carbon and nitrogen reserves and accelerates sink metabolism, leading to improved source-sink relations. The response of plants to defoliation could be used to manipulate source-sink relations by removing lower and senescing leaves to obtain greatest photosynthetic capacity and efficient carbon and nitrogen metabolism under optimal and stressful environments. The present work enhances our current understanding on the physiological responses of plants to defoliation and elaborates how defoliation influences growth, photosynthetic capacity and source-sink relations under optimal and changing environmental conditions.     

Leaf gas exchange of cassava as affected by quality of planting material and water stress

Field trial was conducted to study the effects of quality of planting material and prolonged water stress on leaf gas exchange of the cassava (Manihot esculenta Crantz) cultivar M Col 1684. Nutrient contents of planting material affected rootlet formation, but not leaf gas exchange. Net photosynthetic rate (PN), stomatal conductance (gs), and intercellular CO2 concentration (Ci) were significantly reduced by prolonged water stress. New leaves developed after recovery from water stress showed higher PN and gs, as compared to leaves of similar ages of unstressed plants. The higher PN was associated with higher leaf nutrient contents, indicating that photosynthetic capacity was enhanced in these leaves. These compensating characteristics may partly explain the small yield reduction often observed in stressed cassava.     

The influences of host plant and genetic resistance to Bacillus thuringiensis on trade-offs between offspring number and growth rate in cabbage loopers, Trichoplusia ni

Abstract.  1. Variation in progeny size and quality is common among insects and this variation can strongly influence individual fitness. Larger progeny typically survive better and develop faster under adverse conditions and may have higher fecundity. Due to resource limitations, however, trade-offs may arise between having fewer large offspring or more smaller ones.
2. For cabbage loopers, Trichoplusia ni , pepper leaves are a poorer larval host than cucumber or tomato leaves as indicated by survival, development rate, and body size. Moths reared on cucumber produced more slower growing offspring than those that had been reared on pepper, which produced fewer, faster growing progeny.
3. Traits conferring resistance to Bacillus thuringiensis ( Bt ) generally are associated with strong deleterious effects that may influence resource allocation and reproductive trade-offs between progeny size and number.
4. Unlike the host-plant related trade-off between progeny size and fecundity observed among susceptible control moths, Bt -resistant parents had both the lowest fecundity and smallest progeny size on all host plants. This finding suggests that the progeny size–number relationship is constrained in resistant individuals.     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

Limited scope for maternal effects in aphid defence against parasitoids

Abstract.  1. A mother's environment frequently affects her offspring's phenotype. Such maternal effects may be adaptive, in particular with respect to pathogens or parasites, for example if maternal exposure increases offspring resistance.
2. In aphids, maternal effects are likely to occur as a result of their telescoping generations. This study investigated whether maternal effects influence the susceptibility of the peach-potato aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), to its parasitoid Diaeretiella rapae (M'Intosh) (Hymenoptera: Braconidae: Aphidiinae).
3. In a first experiment, susceptibility was compared among offspring of aphid mothers that had either no contact to parasitoids, had contact but were not attacked, or were attacked but not mummified. Mothers from the last group had successfully resisted the parasitoid.
4. In a second experiment using two different clones, maternal and progeny environment were manipulated by rearing each generation either on a benign (radish) or a more stressful host plant (silver beet) before progeny exposure to parasitoids.
5. The first experiment revealed no significant effect of the maternal treatment on offspring susceptibility to parasitoids and thus no evidence for trans-generational defence. In the second experiment, maternal environment effects were also weak, yet with a trend towards less susceptible offspring of aphid mothers reared on the more stressful plant. However, there was a significant difference among clones and a strong clone × progeny host plant interaction, illustrating that the outcome of a parasitoid attack may be determined by a complex interplay of genetic and environmental factors.
6. Overall, the results suggest that there is limited scope for maternal effects in aphid defence against parasitoids.