Salinity tolerance and genetic variability in freshwater and brackish Iris hexagona colonies

? Premise of the study: Saltwater intrusion is one of the most widespread environmental threats to freshwater wetlands. Iris species worldwide are important members of these plant communities. Wetland irises reproduce clonally and sexually, which permits populations to spread and disperse in benign and stressful conditions. The ability of iris populations to tolerate and adapt to elevated salinity can play an important role in determining the long-term health of wetland ecosystems. ? Methods: We used microsatellite markers to evaluate population structure and genetic diversity, and we performed a common garden experiment to examine the effect of salinity on the growth and reproduction of wild Iris hexagona collected from freshwater and brackish wetlands. ? Key results: Colonies were genetically distinct, with average to high heterozygosity (0.55-0.66) for a clonal species. Salinity had negative linear effects on leaf mass (g), clonal growth (g), root mass (g), and flower numbers, and it had nonlinear effects on seed numbers and seed mass (mg). The greatest sexual reproduction occurred in the intermediate-salinity (4 parts per thousand) treatment. Flowering phenology was delayed for 5 days in the highest-salinity treatment. ? Conclusions: We hypothesized that irises from brackish habitats would tolerate salinity better than freshwater irises would, but no difference in iris performance existed between the two habitats. The observed salinity tolerance and genetic diversity of I. hexagona indicate that populations will persist despite moderate increases in environmental salinity.     

Separate and interacting effects of deer florivory and salinity stress on iris herbivores

Most herbivore studies focus on foliage feeders, despite the potentially strong impact of florivores on plant fitness and ecological communities. Vertebrate florivory is particularly noteworthy because it destroys the inflorescence, and may affect the community of floral arthropods. Strong biotic forces such as florivory can be influenced by abiotic forces. For example, field studies indicate that white-tailed deer Odocoileus virginianus consume up to 90% of all flowers produced in Iris hexagona populations that inhabit high salinity wetlands, whereas freshwater iris populations suffer relatively little damage. We conducted a common garden experiment to investigate the potential effects of vertebrate florivory and salinity on two iris herbivores, the leafminer Cerodontha iridiphora , and the verbena bud moth, Endothenia hebesana . We administered four levels of florivory: zero (control), one, two, and three flowers removed per stalk, and applied three levels of salinity (0, 4 and 8 parts per thousand NaCl) in a full-factorial design. Florivory and salinity had separate and interacting effects on herbivore performance. Salinity was more detrimental to herbivores than florivory, which was beneficial at low levels. A positive effect of moderate salinity on sexual reproduction was reversed by florivory. Our research reveals complex interactions between environmental stress and florivory, an understudied form of herbivory. These interactions have implications for wetland ecological communities that are increasingly exposed to biotic and abiotic stress.     

Flowering phenology in natural populations of Iris pumila

In the present study, I examined flowering phenology of dwarf bearded ins, Iris pumila , using naturally occurring clones (three sites, four microsites) and clones transplanted between two sites representing two habitats Naturally occurring clones in contrasting habitats and microsites differed significantly in phenology, with exposition of the site or microsite being the most important Genotypes from more exposed habitats flowered earlier in both habitats and these differences were statistically significant Patterns of between-habitat and between-population differences were stable over years even though years differed markedly in the flowering onset Within-population genetic variability for flowering phenology was also found to be significant Although clones with dark-colored flowers tended to flower earlier in all habitats and microsites I failed to detect statistically significant differences in flowering time among color morphs Flowering phenology in I pumila is highly susceptible to environmental variability, but this factor has not prevented population divergence in flowering time Between-habitat differences in flowering time turned out to be a result of both phenotypic plasticity of individuals and genetic differentiation of populations     

The deleterious effects of salinity stress on leafminers and their freshwater host

Abstract.  1. Salinity is an important cause of abiotic stress in wetland communities yet little is known about its consequences for freshwater plants and their insect herbivores. The goal of this study was to test the effect of salinity on a leafmining insect, Cerodontha iridiphora , and its herbaceous host plant, Iris hexagona .
2. Leafminer performance was evaluated on irises grown in control and saline treatments, and the effects of salinity and herbivory on leaf quality and mortality was measured.
3. Leafminer density and size were significantly lower on irises grown in saline water compared with freshwater.
4. Both salinity and herbivory accelerated leaf senescence and mortality, and their combined effects increased tissue loss by an order of magnitude compared with controls.
5. Leafminers acted as nutrient sinks. The undamaged regions of mined leaves contained 40% less nitrogen than unmined leaves, providing a mechanism for the premature leaf mortality.
6. Salinity was detrimental to the performance and survival of both the iris leafminer and its host plant. We propose that glycophytic host plants and their insect herbivores will suffer more than halophytic communities from environmental salinity because they lack the adaptive mechanisms to tolerate this potent physiological stress.     

迁地保育植物花期物候对迁入地气候变化的响应

以中国科学院武汉植物园内栽培的长果秤锤树(Sinojackia dolichocarpa C. J. Qi)、山白树(Sinowilsonia henryi Hemsl.)、夏腊梅(Sinocalycanthus chinensis Cheng et S. Y. Chang)、紫茎(Stewartia sinensis Rehd. et Wils.)和绒毛皂荚(Gleditsia vestita Chun et How ex B. G. Li) 5种迁地保育植物为对象,通过2008-2016年观察记录的初花期物候及整个花期长度的数据,研究花期的年际变化规律及其与迁入地武汉气候因子的相关性。结果显示:(1)从初花期来看,长果秤锤树的初花期每年提前1.25 d,紫茎的初花期每年推迟1.35 d,绒毛皂荚的初花期每年推迟1.22 d。(2)从花期长度来看,山白树的花期每年增加1.72 d,夏蜡梅的花期每年减少1.62 d,紫茎的花期每年增加0.32 d。(3)从花期与气候因子的相关性来看,年降水量、年平均相对湿度、 10℃有效积温、花前 10℃的有效积温是影响这5种植物初花期、花期长度的主要气候因子;不同物种间影响花期的主要气候因子有所差异。     

Flowering phenology and compensation for herbivory in<Emphasis Type="Italic"> Ipomopsis aggregata</Emphasis>

The mechanisms and circumstances that affect a plant's ability to tolerate herbivory are subjects of ongoing interest and investigation. Phenological differences, and the timing of flowering with respect to pollinators and pre-dispersal seed predators, may provide one mechanism underlying variable responses of plants to herbivore damage. The subalpine wildflower, Ipomopsis aggregata, grows across a wide range of elevations and, because phenology varies with elevation, phenological delays associated with elevation may affect the ability of I. aggregata to compensate for or tolerate browsing. Thus, we examined the response of I. aggregata to herbivory across an elevation gradient and addressed the interactions among phenological delays imposed by damage, elevation, pre-dispersal seed predation and pollination, on I. aggregata's compensatory response. Among high and low elevation populations in areas near the Rocky Mountain Biological Laboratory (RMBL) in Gothic, Colorado, we compared the responses of naturally browsed, artificially browsed (clipped), and unbrowsed (control) plants of I. aggregata. We compared responses in the date of initiation of flowering, timing of peak bloom, floral display, nectar production and sugar concentration, oviposition and fruit destruction by the pre-dispersal seed predator Hylemya sp. (Anthomyiidae), fruit production, and aboveground biomass production. Clipping had the greatest effect on reproductive success and clipped plants at high elevation exhibited the lowest tolerance for herbivory. The effects of browsing appear to be mediated by flowering phenology, and both browsing and elevation delayed flowering phenology. Time needed for regrowth delays flowering, and thus affects the overlap with seed predators and pollinators. As a result of delayed flowering, naturally browsed and clipped plants incurred lower rates of seed predation. In the absence of seed predation, plants would exhibit a lower tolerance to herbivory since naturally and artificially browsed plants had fewer fruits destroyed by Hylemya larvae. We provide additional evidence that, for populations near the RMBL, clipping and natural browsing do not have the same effect on I. aggregata plants. This may be due to the selection of larger plants by herbivores. Although under some conditions plants may tolerate browsing, in areas where the growing season is short a phenological delay imposed by damage is likely to significantly reduce plant fitness. Identifying the mechanisms that allow plants to tolerate herbivore damage will help to develop a general framework for understanding the role of tolerance in plant population and community dynamics, as well as plant-herbivore interactions.     

外源水杨酸改善Cd胁迫下路易斯安娜鸢尾的生长和养分吸收

采用Hoagland溶液培养,研究了外源水杨酸(SA)预处理对镉(Cd)胁迫下路易斯安娜鸢尾生物量、含水量、光合作用、根系活力、Cd积累、矿质养分吸收和抗氧化酶活性的影响.结果表明： Cd胁迫下,SA预处理的植株根叶干质量、根系活力和净光合速率提高.与单独Cd处理相比,SA预处理后的胁迫植株Cd含量无明显变化,而叶片N、P、S含量显著提高,K含量显著下降,根部抗氧化酶活性显著提高.表明SA预处理可明显缓解路易斯安娜鸢尾的Cd伤害,其原因不在于减少Cd吸收量而是提高了对Cd的抗性,矿质元素的吸收积累调节和根部抗氧化酶活性的提高是其毒害缓解的重要原因.     

Informing native plant sourcing for ecological restoration: cold‐hardiness dynamics,flowering phenology,and survival of Eriogonum umbellatum

Despite advances in restoration of degraded lands around the world, native plants are still underutilized. Selection of appropriate plant materials is a critical factor in determining plant establishment and persistence. To better inform decision‐making, we examined cold‐hardiness dynamics, flowering phenology, and survival among five geographically distinct sulfur‐flower buckwheat (Polygonaceae: Eriogonum umbellatum Torr.) populations in a common garden. LT₅₀ (a measure of freezing injury) was determined every 6 weeks across a complete year; one population was also evaluated at the source. Cold‐hardiness dynamics were similar across populations, with annual fluctuations in mean LT₅₀ exceeding 40°C. Rate of deacclimation (i.e. loss of cold tolerance) in spring varied across populations and was not related to the elevation from which a population came. Plants were less cold hardy in October 2014 compared to October 2013, likely reflecting a response to colder local conditions in 2013. Although the range of LT₅₀ was similar for a single comparison of common garden versus wild‐grown plants, wild‐grown plants acclimated and deacclimated earlier than common garden‐grown plants. Plants derived from a low‐elevation population showed delayed flowering phenology, while high‐elevation populations showed earlier flowering phenology, with one high‐elevation population having the lowest survival rate in the common garden. These results suggest that while considerable plasticity in seasonal cold‐hardiness dynamics occur, population variability in deacclimation and flowering phenology have implications for selection and movement of sulfur‐flower buckwheat for ecological restoration.     

Selection on flowering time and floral display in an alpine and a lowland population of Arabidopsis lyrata

To determine whether population differentiation in flowering time is consistent with differences in current selection, we quantified phenotypic selection acting through female reproductive success on flowering phenology and floral display in two Scandinavian populations of the outcrossing, perennial herb Arabidopsis lyrata in two years. One population was located in an alpine environment strongly affected by grazing, whereas the other was close to sea level and only moderately affected by herbivory. Multiple regression models indicated directional selection for early end of flowering in one year in the lowland population, and directional selection for early start of flowering in one year in the alpine population. As expected, there was selection for more inflorescences in the lowland population. However, in the alpine population, plants with many inflorescences were selectively grazed and the number of inflorescences produced was negatively related to female fitness in one year and not significantly related to female fitness in the second year. The results are consistent with the hypothesis that genetic differentiation in flowering phenology between the study populations is adaptive, and indicate that interactions with selective grazers may strongly influence selection on floral display in A. lyrata.     

Ontogeny of flower parts on naturally growing Iris pumila clones: implications for population differentiation and phenotypic plasticity studies

Previous studies revealed significant phenotypic plasticity, genetic variability and population differentiation of flower morphometric traits on dwarf bearded iris Iris pumila. Also, study of I. pumila flowering phenology revealed significant impact of habitat type as well as population differentiation for flowering time. Since the flowering time can influence other flower traits, we performed this analysis of flower morphometric traits in three time points during the flower bud ontogenic development in two habitat types (open vs. shaded). Analysis revealed that for most of the traits greater trait values were recorded for open habitat but only on latter time points. For most of the analyzed traits direction of differences in bud stage was the opposite to the direction of differences in mature flower stage detected in previous studies. However, length of the stem, a trait that showed the greatest variability between habitats and populations and therefore greatest genetic differentiation and phenotypic plasticity, was significantly greater in the samples from the late flowering shaded habitat in all time samples, indicating that in case of this trait different mechanisms were involved. Those findings have implications for design of the future studies on I. pumila.     

冬季水淹对秋华柳的开花物候及繁殖分配的影响

开花物候及繁殖分配是植物适应环境的重要因素,为了解长期冬季水淹对三峡库区耐淹物种秋华柳(Salix variegata Franch.)繁殖的影响,研究了长期冬季水淹条件下秋华柳的开花物候和繁殖分配情况。实验在2006年11月份设置了如下处理:对照,完全水淹(植株置于水中,顶部距水面2m)30,60,90,120d和150d。结果表明:(1)对照及各水淹处理的秋华柳花期都较长,在7-11月份持续开花,个体开花进程(开花振幅曲线)呈单峰曲线。(2)冬季水淹对秋华柳群体及个体的开花物候有显著影响。水淹时间越长,始花期越晚,花期持续时间越短(P0.05)。(3)长期冬季水淹下,秋华柳显著降低了繁殖分配比例和全株生物量及单株花序数(P0.05)。(4)开花物候指数与繁殖分配的相关分析表明:始花时间越晚的个体,花期持续时间越短。花期持续时间越短的个体花序数越少,致使繁殖分配越小。总的来说,冬季水淹下,秋华柳通过推迟开花日期、缩短花期持续时间使繁殖分配比例降低,将更多的资源分配到生存力上,是秋华柳对长期冬季水淹的一种适应。同时,在长期冬季水淹后,秋华柳仍保持一定的开花繁殖能力,是其在应用于三峡水库消落区植被构建后产生后代延续种群的前提条件。     

Site-to-site differences in pollinator visitation patterns in a Louisiana iris hybrid zone

We studied pollinator visitation rates and movement patterns in experimental arrays of irises at two sites within a Louisiana iris hybrid zone. Arrays contained single-flowered stems of red-flowered I. fulva , blue-flowered I. hexagona , and purple-flowered F1 hybrids. At one site, where I. hexagona was the only wild-growing iris, queen bumble bees were the most common pollinator, and the rank order of pollinator visit rates was I. hexagona I. fulva . At a second site, where I. fulva predominated in the wild, hummingbirds were the most common pollinators, and this order was reversed: I. hexagona 相似文献   

Response to joint selection on germination and flowering phenology depends on the direction of selection

Flowering and germination time are components of phenology, a complex phenotype that incorporates a number of traits. In natural populations, selection is likely to occur on multiple components of phenology at once. However, we have little knowledge of how joint selection on several phenological traits influences evolutionary response. We conducted one generation of artificial selection for all combinations of early and late germination and flowering on replicated lines within two independent base populations in the herb Campanula americana. We then measured response to selection and realized heritability for each trait. Response to selection and heritability were greater for flowering time than germination time, indicating greater evolutionary potential of this trait. Selection for earlier phenology, both flowering and germination, did not depend on the direction of selection on the other trait, whereas response to selection to delay germination and flowering was greater when selection on the other trait was in the opposite direction (e.g., early germination and late flowering), indicating a negative genetic correlation between the traits. Therefore, the extent to which correlations shaped response to selection depended on the direction of selection. Furthermore, the genetic correlation between timing of germination and flowering varies across the trait distributions. The negative correlation between germination and flowering time found when selecting for delayed phenology follows theoretical predictions of constraint for traits that jointly determine life history schedule. In contrast, the lack of constraint found when selecting for an accelerated phenology suggests a reduction of the covariance due to strong selection favoring earlier flowering and a shorter life cycle. This genetic architecture, in turn, will facilitate further evolution of the early phenology often favored in warm climates.     

Olive phenology as a sensitive indicator of future climatic warming in the Mediterranean

Experimental and modelling work suggests a strong dependence of olive flowering date on spring temperatures. Since airborne pollen concentrations reflect the flowering phenology of olive populations within a radius of 50 km, they may be a sensitive regional indicator of climatic warming. We assessed this potential sensitivity with phenology models fitted to flowering dates inferred from maximum airborne pollen data. Of four models tested, a thermal time model gave the best fit for Montpellier, France, and was the most effective at the regional scale, providing reasonable predictions for 10 sites in the western Mediterranean. This model was forced with replicated future temperature simulations for the western Mediterranean from a coupled ocean‐atmosphere general circulation model (GCM). The GCM temperatures rose by 4·5 °C between 1990 and 2099 with a 1% per year increase in greenhouse gases, and modelled flowering date advanced at a rate of 6·2 d per °C. The results indicated that this long‐term regional trend in phenology might be statistically significant as early as 2030, but with marked spatial variation in magnitude, with the calculated flowering date between the 1990s and 2030s advancing by 3–23 d. Future monitoring of airborne olive pollen may therefore provide an early biological indicator of climatic warming in the Mediterranean.     

Increased variance in temperature and lag effects alter phenological responses to rapid warming in a subarctic plant community

Summer temperature on the Cape Churchill Peninsula (Manitoba, Canada) has increased rapidly over the past 75 years, and flowering phenology of the plant community is advanced in years with warmer temperatures (higher cumulative growing degree days). Despite this, there has been no overall shift in flowering phenology over this period. However, climate change has also resulted in increased interannual variation in temperature; if relationships between phenology and temperature are not linear, an increase in temperature variance may interact with an increase in the mean to alter how community phenology changes over time. In our system, the relationship between phenology and temperature was log‐linear, resulting in a steeper slope at the cold end of the temperature spectrum than at the warm end. Because below‐average temperatures had a greater impact on phenology than above‐average temperatures, the long‐term advance in phenology was reduced. In addition, flowering phenology in a given year was delayed if summer temperatures were high the previous year or 2 years earlier (lag effects), further reducing the expected advance over time. Phenology of early‐flowering plants was negatively affected only by temperatures in the previous year, and that of late‐flowering plants primarily by temperatures 2 years earlier. Subarctic plants develop leaf primordia one or more years prior to flowering (preformation); these results suggest that temperature affects the development of flower primordia during this preformation period. Together, increased variance in temperature and lag effects interacted with a changing mean to reduce the expected phenological advance by 94%, a magnitude large enough to account for our inability to detect a significant advance over time. We conclude that changes in temperature variability and lag effects can alter trends in plant responses to a warming climate and that predictions for changes in plant phenology under future warming scenarios should incorporate such effects.     

Ontogeny of flower parts on naturally growing <Emphasis Type="Italic">Iris pumila</Emphasis> clones: Implications for population differentiation and phenotypic plasticity studies

Previous studies revealed significant phenotypic plasticity, genetic variability and population differentiation of flower morphometric traits on dwarf bearded iris Iris pumila. Also, study of I. pumila flowering phenology revealed significant impact of habitat type as well as population differentiation for flowering time. Since the flowering time can influence other flower traits, we performed this analysis of flower morphometric traits in three time points during the flower bud ontogenic development in two habitat types (open vs. shaded). Analysis revealed that for most of the traits greater trait values were recorded for open habitat but only on latter time points. For most of the analyzed traits direction of differences in bud stage was the opposite to the direction of differences in mature flower stage detected in previous studies. However, length of the stem, a trait that showed the greatest variability between habitats and populations and therefore greatest genetic differentiation and phenotypic plasticity, was significantly greater in the samples from the late flowering shaded habitat in all time samples, indicating that in case of this trait different mechanisms were involved. Those findings have implications for design of the future studies on I. pumila.     

Selection for phenotypic divergence between diploid and autotetraploid Heuchera grossulariifolia

Much of the diversity of flowering plants is associated with genomic duplication through polyploidy. Little is known, however, about the evolutionary mechanisms responsible for the diversification of novel polyploid lineages. We evaluated the possibility that divergence is driven by natural selection by estimating the strength of phenotypic selection acting on three floral traits in sympatric populations of diploid and autotetraploid Heuchera grossulariifolia over three years. Our results demonstrate consistent directional selection for increasing scape length and floral display in both diploid and tetraploid populations. In contrast, selection acting on flowering phenology varied across year and ploidy. Specifically, selection was found to favor late-flowering diploids in 2001 and 2002 but early-flowering tetraploids in 2003. We investigated the mechanistic basis of divergent selection for flowering phenology in 2003 by estimating the relationship between plant flowering phenology and the probability of intercytotype pollinator movement. The results demonstrated that less divergent tetraploids were significantly more likely to experience intercytotype flights than were more divergent tetraploids. This result is consistent with the pattern of phenotypic selection observed. Taken together, our results suggest that divergence of polyploids and their diploid progenitors may be driven by a process analogous to reinforcement whereby selection favors phenotypes that reduce the probability of intercytotype matings with reduced fertility.     

Interactive Effects of Arbuscular Mycorrhizal Fungi and Copper Stress on Flowering Phenology and Reproduction of Elsholtzia splendens

Plant responses to heavy metal contamination may depend on the presence of arbuscular mycorrhizal fungi (AMF). Elsholtzia splendens is an indicator species for the presence of copper (Cu) mines because both its flowering phenology and reproduction are tolerant to heavy metals. To test whether effects of Cu on the flowering phenology and reproduction of E. splendens depend on the presence of AMF, we conducted a factorial experiment with two Cu treatments (with or without Cu addition) crossed with two AMF treatments (with or without AMF inoculation). Without AMF, Cu addition significantly delayed the onset dates, ending dates and peak dates of flowering and decreased flowering duration. However, AMF inoculation reversed the effects of Cu stress, with recovered flowering onset and ending dates and increased the flowering duration. Cu addition significantly decreased inflorescence width and number, inflorescence biomass, vegetative biomass and total seed number, but significantly increased 1000-seed weight. AMF inoculation significantly increased vegetative biomass. Two-way ANOVA results showed that the interactive effects between Cu addition and AMF inoculation were significant on the inflorescence number, vegetative biomass and total seed number. These results indicate that AMF can alleviate the Cu stress on the flowering phenology and reproduction of E. splendens.     

Response to simulated climatic change in an alpine and subarctic pollen-risk strategist, Silene acaulis

The aim of this study was to test if early flowering species respond with increased seed production to climate warming as is predicted for late-flowering seed-risk strategists. Experimental climate warming of about 3°C was applied to two populations of the cushion-forming plant Silene acaulis (L.) Jacq. The experiment was run at one subarctic site and one alpine site for 2 years and 1 year, respectively, using open-top chambers (OTC).
The 2-year temperature enhancement at the subarctic site had a marked effect on the flowering phenology. Cushions inside the OTC started flowering substantially earlier than control cushions. Both the male and female phases developed faster in the OTCs, and maturation of capsules occurred earlier. The cushions also responded positively in reproductive terms and produced more mature seeds and had a higher seed/ovule ratio. After 1 year temperature enhancement at the alpine site there was a weak trend for earlier flowering, but there was no significant difference in seed production or seed/ovule ratio.