0000000000037371
AUTHOR
Sandra Varga
Sex-specific interaction between arbuscular mycorrhizal and dark septate fungi in the dioecious plant Antennaria dioica (Asteraceae).
Male and female plants of dioecious species often differ in their resource demands and this has been linked to secondary sexual dimorphism, including sex-specific interactions with other organisms such as herbivores and pollinators. However, little is known about the interaction between dioecious plants and fungal root endophytes. Plants may be simultaneously colonised by arbuscular mycorrhizal (AM) and dark septate (DS) fungi. While it is well established that AM mutualism involves reciprocal transfer of photosynthates and mineral nutrients between roots of host plants and these fungi, the role of DS fungi remains controversial. Here, we report the temporal and spatial variation in AM and …
Prolonged stigma and flower lifespan in females of the gynodioecious plant Geranium sylvaticum
tIn gynodioecious plants females need a reproductive advantage over hermaphrodites to be maintainedin the same population. Generally, three main proximate causes for a female advantage are considered:inbreeding avoidance, different resource allocation patterns, and differences in ecological interactions.A mechanism potentially causing a female advantage that is rarely discussed is a difference in flo-ral longevity between the genders. Females may have a longer stigma lifespan than hermaphrodites,which can affect pollination. Stigma and flower lifespan are rarely documented in gynodioecious species,although it is a common observation in dioecious species that female plants flower longer than…
Sex-specific patterns of antagonistic and mutualistic biotic interactions in dioecious and gynodioecious plants
a b s t r a c t A major transition in flowering plants has been the evolution of separate sexes from hermaphroditism via gynodioecy which is considered to be the most important route. Biotic interactions, both antagonist and mutualistic, have been proposed to influence this transition which is generally accompanied by the evolution of sexual dimorphism in secondary sexual traits. While some researchers have studied sex- specific patterns in herbivory and pollination, less attention has been paid to pathogens/parasites and a limited number of studies have revised sex-specific patterns in mycorrhizal symbiosis. In this article, we explore sex-specific interactions in dioecious and gynodioecio…
Prolonged stigma and flower lifespan in females of the gynodioecious plant Geranium sylvaticum
In gynodioecious plants females need a reproductive advantage over hermaphrodites to be maintained in the same population. Generally, three main proximate causes for a female advantage are considered: inbreeding avoidance, different resource allocation patterns, and differences in ecological interactions. A mechanism potentially causing a female advantage that is rarely discussed is a difference in floral longevity between the genders. Females may have a longer stigma lifespan than hermaphrodites, which can affect pollination. Stigma and flower lifespan are rarely documented in gynodioecious species, although it is a common observation in dioecious species that female plants flower longer t…
Sex-specific responses to mycorrhiza in a dioecious species.
In most studies about dioecious plants, the role of arbuscular mycorrhizae (AM) and the potential sex-specific differences between the plant hosts have been overlooked. Because plant sexes frequently differ in drought tolerance and AM fungal colonization provides higher resistance to drought, we investigated whether the relation of mycorrhizal fungi with either male or female Antennaria dioica plants differs using a factorial experiment. We hypothesized that because AM usually increase growth rate and male plants usually grow larger than females, males should gain more benefit from the mycorrhizal symbiosis in terms of mineral nutrition and water supply. Because of higher demands of carbohy…
Interrelationships between mycorrhizal symbiosis, soil pH and plant sex modify the performance of Antennaria dioica
AM symbiosis is usually beneficial for plants, but the benefits gained may depend on the soil abiotic factors. In dioecious plants, female and male individuals have different resource demands and allocation patterns. As a consequence of these differences, it is logical to assume that female and male plants differ in their relationship with arbuscular mycorrhizal (AM) fungi, although this has rarely been examined. We used a factorial greenhouse experiment to investigate whether female and male plants in the dioecious model species Antennaria dioica have a different relationship with their AM symbionts under two soil pH levels. In particular, we asked: (1) Do the sexes in A. dioica have sex-s…
On the importance of details in arbuscular mycorrhizal research
Most journals require authors to provide sufficient experimental detail in their publications to enable other scientists to reproduce the studies presented. However, my personal experience when reading papers in my research field suggests that many details that could be considered important are commonly overlooked. I analysed the work published during 2013 within the field of arbuscular mycorrhizal ecology, assessing whether 15 important details from 5 fundamental criteria were reported about (1) the experimental treatment, (2) the abiotic growing conditions, (3) the soil nutrient concentrations, (4) the duration of the study and (5) a description of the methodology employed to collect the …
Competitive interactions are mediated in a sex-specific manner by arbuscular mycorrhiza in Antennaria dioica
Plants usually interact with other plants, and the outcome of such interaction ranges from facilitation to competition depending on the identity of the plants, including their sexual expression. Arbuscular mycorrhizal (AM) fungi have been shown to modify competitive interactions in plants. However, few studies have evaluated how AM fungi influence plant intraspecific and interspecific interactions in dioecious species. The competitive abilities of female and male plants of Antennaria dioica were examined in a greenhouse experiment. Females and males were grown in the following competitive settings: (i) without competition, (ii) with intrasexual competition, (iii) with intersexual competitio…
Sexes in gynodioecious Geranium sylvaticum do not differ in their isotopic signature or photosynthetic capacity
In gynodioecious plants, females are expected to produce more or better seeds than hermaphrodites in order to be maintained within the same population. Even though rarely measured, higher seed production can be achieved through differences in physiology. In this work, we measured sexual dimorphism in several physiological traits in the gynodioecious plant Geranium sylvaticum. Photosynthetic rate, stomatal conductance, transpiration rate, WUE and isotopic signatures were measured in plants growing in two habitats differing in light availability. Females have been reported to produce more seeds than hermaphrodites. However, we did not observe any significant difference in seed output between …
Absence of Sex Differential Plasticity to Light Availability during Seed Maturation in Geranium sylvaticum
Sex-differential plasticity (SDP) hypothesis suggests that since hermaphrodites gain fitness through both pollen and seed production they may have evolved a higher degree of plasticity in their reproductive strategy compared to females which achieve fitness only through seed production. SDP may explain the difference in seed production observed between sexes in gynodioecious species in response to resource (nutrients or water) availability. In harsh environments, hermaphrodites decrease seed production whereas females keep it relatively similar regardless of the environmental conditions. Light availability can be also a limiting resource and thus could theoretically affect differently femal…
Dioecious species and arbuscular mycorrhizal symbioses: the case of Antennaria dioica.
Sex-specific interactions with herbivores and pollinators have been observed in female and male plants of dioecious species. However, only a limited number of studies have revised sex-specific patterns in mycorrhizal symbiosis. To test whether female and male plants of Antennaria dioica differ in their relationship with arbuscular mycorrhizal (AM) fungi, we examined the temporal and spatial variation in AM fungi in female, male and non-reproductive A. dioica plants in three natural populations in Finland during flowering and after seed production. Our results are consistent with previous studies both under greenhouse and field conditions with the same species showing differences in AM colon…
Sexes in gynodioeciousGeranium sylvaticumdo not differ in their isotopic signature or photosynthetic capacity
• In gynodioecious plants, females are expected to produce more or better seeds than hermaphrodites in order to be maintained within the same population. Even though rarely measured, higher seed production can be achieved through differences in physiology. • In this work, we measured sexual dimorphism in several physiological traits in the gynodioecious plant Geranium sylvaticum. Photosynthetic rate, stomatal conductivity, transpiration rate, water use efficiency and isotopic signatures were measured in plants growing in two habitats differing in light availability. • Females have been reported to produce more seeds than hermaphrodites. However, we did not observe any significant difference…
Faster acquisition of symbiotic partner by common mycorrhizal networks in early plant life stage
Arbuscular mycorrhizal (AM) fungi usually improve plant performance yet our knowledge about their effects on seed germination and early plant establishment is very limited. We performed a factorial greenhouse experiment where the seeds from four low Arctic cooccurring mycorrhizal herbs (Antennaria dioica, Campanula rotundifolia, Sibbaldia procumbens, and Solidago virgaurea) were germinated alone or in the vicinity of an adult Sibbaldia plant with or without AM fungi; given either as spores or being present in a common mycorrhizal network (CMN). Three different AM fungal species were examined to assess species-specific differences in symbiont acquisition rate. Out of the four plant species i…
Host plant and arbuscular mycorrhizal fungi show contrasting responses to temperature increase: Implications for dioecious plants
Individual plants live in complex environments where they interact with other organisms such as herbivores, pollinators, fungi and pathogens. The influence of rising temperature on biotic interactions has begun to receive attention, and is an important research frontier currently. However, the belowground interactions with organisms such as arbuscular mycorrhizal (AM) fungi have received little attention so far. In this study, we investigated the response of the dioecious plant Antennaria dioica and its AM fungi to increased temperature in a controlled environment simulating the period of growth of A. dioica in central Finland. Specifically, we evaluated the effect of rising temperature on …
Variable mycorrhizal benefits on the reproductive output ofGeranium sylvaticum, with special emphasis on the intermediate phenotype
In several gynodioecious species, intermediate sex between female and hermaphrodite has been reported, but few studies have investigated fitness parameters of this intermediate phenotype. Here, we examined the interactions between plant sex and arbuscular mycorrhizal (AM) fungal species affecting the reproductive output of Geranium sylvaticum, a sexually polymorphic plant species with frequent intermediate sexes between females and hermaphrodites, using a common garden experiment. Flowering phenology, AM colonisation levels and several plant vegetative and reproductive parameters, including seed and pollen production, were measured. Differences among sexes were detected in flowering, fruit …
Effects of arbuscular mycorrhizal fungi and maternal plant sex on seed germination and early plant establishment.
Premise of the study: Arbuscular mycorrhizal fungi usually enhance overall plant performance, yet their effects on seed germi- nation and early plant establishment, crucial steps in plant cycles, are generally overlooked. In gynodioecious species, sexual dimorphism in these traits has been reported, with females producing seeds that germinate at a faster rate than seeds from hermaphrodites. • Methods: Using the gynodioecious plant Geranium sylvaticum, I investigated in a greenhouse experiment whether the presence of arbuscular mycorrhizal spores affects seed germination and early plant establishment, examining at the same time whether the sex of the mother producing the seeds also influence…
Gender dimorphism and mycorrhizal symbiosis affect floral visitors and reproductive output in Geranium sylvaticum
1. Arbuscular mycorrhizal (AM) symbiosis has been shown to enhance some plant traits to which pollinators are known to respond. Moreover, in gynodioecious species pollinators prefer hermaphrodite flowers over female ones, but the role of fungal symbiosis in sex-specific pollinator attraction is unknown. 2. We examined how plant gender dimorphism and AM symbiosis affect floral visitors and reproductive output in the gynodioecious plant Geranium sylvaticum. Floral visitors were moni- tored in a common garden experiment using mycorrhizal plants inoculated with either Glomus claroideum or Glomus hoi and in non-mycorrhizal condition. 3. We hypothesized that because of the larger flower display a…
Sex ratio and spatial distribution of male and female Antennaria dioica (Asteraceae) plants
Sex ratio, sex spatial distribution and sexual dimorphism in reproduction and arbuscular mycorrhizal colonisation were investigated in the dioecious clonal plant Antennaria dioica (Asteraceae). Plants were monitored for five consecutive years in six study plots in Oulanka, northern Finland. Sex ratio, spatial distribution of sexes, flowering frequency, number of floral shoots and the number and weight of inflorescences were recorded. In addition, intensity of mycorrhizal fungi in the roots was assessed. Both sexes flowered each year with a similar frequency, but the overall genet sex ratio was strongly female-biased. The bivariate Ripley’s analysis of the sex distribution showed that within…
Nectar sugar production across floral phases in the Gynodioecious Protandrous Plant Geranium sylvaticum [corrected].
Many zoophilous plants attract their pollinators by offering nectar as a reward. In gynodioecious plants (i.e. populations are composed of female and hermaphrodite individuals) nectar production has been repeatedly reported to be larger in hermaphrodite compared to female flowers even though nectar production across the different floral phases in dichogamous plants (i.e. plants with time separation of pollen dispersal and stigma receptivity) has rarely been examined. In this study, sugar production in nectar standing crop and secretion rate were investigated in Geranium sylvaticum, a gynodioecious plant species with protandry (i.e. with hermaphrodite flowers releasing their pollen before th…
Differential competitive ability between sexes in the dioecious Antennaria dioica (Asteraceae)
† Background and Aims Differences in competitive ability between the sexes of dioecious plants are expected as a result of allocation trade-offs associated with sex-differential reproductive costs. However, the available data on competitive ability in dioecious plants are scarce and contradictory. In this study sexual competition was eval- uated using the dioecious plant Antennaria dioica in a common garden transplantation experiment. † Methods Male and female plants were grown for 3 years either in isolation, or in competition with a plant of the same sex or the opposite sex. Flowering phenology, sexual and asexual reproduction, plant growth, nutrient content and arbuscular mycorrhizal col…
Competitive interactions are mediated in a sex-specific manner by arbuscular mycorrhiza inAntennaria dioica
Plants usually interact with other plants, and the outcome of such interaction ranges from facilitation to competition depending on the identity of the plants, including their sexual expression. Arbuscular mycorrhizal (AM) fungi have been shown to modify competitive interactions in plants. However, few studies have evaluated how AM fungi influence plant intraspecific and interspecific interactions in dioecious species. The competitive abilities of female and male plants of Antennaria dioica were examined in a greenhouse experiment. Females and males were grown in the following competitive settings: (i) without competition, (ii) with intrasexual competition, (iii) with intersexual competitio…
Light availability affects sex lability in a gynodioecious plant.
PREMISE OF THE STUDY: Sex lability (i.e., gender diphasy) in plants is classically linked to the larger resource needs associated with the female sexual function (i.e., seed production) compared to the male function (i.e., pollen production). Sex lability in response to the environment is extensively documented in dioecious species, but has been largely overlooked in gynodioecious plants. METHODS: Here, we tested whether environmental conditions induce sex lability in the gynodioecious Geranium sylvaticum. We conducted a transplantation experiment in the field where plants with different sex expression were reciprocally transplanted between high light and low light habitats. We measured pla…
Mycorrhizal benefit differs among the sexes in a gynodioecious species
Both plant sex and arbuscular mycorrhizal (AM) symbiosis influence resource acquisition and allocation in plants, but the interaction between these two components is not well established. As the different plant sexes differ in their resource needs and allocation patterns, it is logical to presume that they might differ in their relationship with AM as well. We investigate whether the association with AM symbiosis is different according to the host plant sex in the gynodioecious Geranium sylvaticum, of which, besides female and hermaphrodite plants, intermediate plants are also recognized. Specifically, we examine the effects of two different AM fungi in plant mass allocation and phosphorus …
Transgenerational effects of plant sex and arbuscular mycorrhizal symbiosis
In gynodioecious plants, females are predicted to produce more and/or better offspring than hermaphrodites in order to be maintained in the same population. In the field, the roots of both sexes are usually colonized by arbuscular mycorrhizal (AM) fungi. Transgenerational effects of mycorrhizal symbiosis are largely unknown, although theoretically expected. We examined the maternal and paternal effects of AM fungal symbiosis and host sex on seed production and posterior seedling performance in Geranium sylvaticum, a gynodioe- cious plant. We hand-pollinated cloned females and hermaphrodites in symbiosis with AM fungi or in nonmycorrhizal conditions and measured seed number and mass, and see…