Amphicarpic plants: definition,ecology, geographic distribution,systematics, life history,evolution and use in agriculture

Although most plants produce all of their fruits (seeds) aboveground, amphicarpic species produce fruits (seeds) both above‐ and belowground. Our primary aims were to determine the number of reported amphicarpic species and their taxonomic, geographic, life form and phylogenetic distribution, to evaluate differences in the life history of plants derived from aerial and subterranean seeds, to discuss the ecological and evolutionary significance of amphicarpy, to explore the use of amphicarpic plants in agriculture, and to suggest future research directions for studies on amphicarpy. Amphicarpy occurs in at least 67 herbaceous species (31 in Fabaceae) in 39 genera and 13 families of angiosperms distributed in various geographical regions of the world and in various habitats. Seeds from aerial and subterranean fruits differ in size/mass, degree of dormancy, dispersal and ability to form a persistent seed bank, with aerial seeds generally being smaller, more dormant and more likely to be dispersed and to form a seed bank than subterranean seeds. In addition, plants produced by aerial and subterranean seeds may differ in survival and growth, competitive ability and biomass allocation to reproduction. Amphicarpic plants may exhibit a high degree of plasticity during reproduction. Subterranean fruits are usually formed earlier than aerial ones, and plants may produce only subterranean propagules under stressful environmental conditions. Differences in the life histories of plants from aerial and subterranean seeds may be an adaptive bet‐hedging strategy.     

QUANTITATIVE VARIATION OF LIFE HISTORY TRAITS IN AMPHICARPIC PEANUTGRASS (AMPHICARPUM PURSHII) AND ITS EVOLUTIONARY SIGNIFICANCE

Peanutgrass, Amphicarpum purshii, is a predominantly selling annual that exhibits an amphicarpic reproductive strategy, producing spikelets (and seeds) both above and below the soil surface. Both aerial and subterranean spikelets are self-fertile, but only the aerial spikelets are chasmogamous and capable of cross-pollination. This species mostly grows in disturbed areas of the New Jersey Pine Barrens. To examine life history variation, 60 quantitative characters were measured on twelve families raised from seeds collected from the aerial panicles of twelve parents grown in the greenhouse. Only 19 (= 31.7%) of the traits showed significant between-family phenotypic variation, and the mean percentage of the total variation due to family was 14.9 (N = 60 characters). Percentage biomass allocation and vegetative characters showed the highest narrow-sense heritabilities. Traits directly relevant to fitness such as subterranean seed set and seed weight of both seed types showed the least genetic variation. This low variation could reflect genetic fixation and/or developmental canalization of those fitness components important to survival in disturbed, early-successional habitats.     

A modular approach to biomass allocation in an invasive annual (Microstegium vimineum; Poaceae)

The invasive grass Microstegium vimineum grows in low light beneath the canopy of eastern forests in North America by reiteration of modules (phytomers) along a tiller. Basal phytomers are vegetative; terminal phytomers produce a raceme of chasmogamous (CH) spikelets plus an axillary raceme of cleistogamous (CL) spikelets. Additional subterminal phytomers with CL racemes mature basipetally. Allocation to culms, leaves, and CH and CL within phytomers was examined in relation to light conditions for a population in New Jersey, USA. Plants were reared in a greenhouse from seed families of parents in deep shade (2-8% full sun) or sunny, edge habitats. Primary tillers were subdivided into phytomers, dried, and weighed. Tillers from field habitats were similarly treated. For vegetative and subterminal phytomers, allocation to leaves and CH was greatest for the shady habitat. CL allocation decreased from terminal to reproductively immature subterminal phytomers. CH and CL mass was positively correlated with leaf mass, suggesting that reproduction is determined by available photosynthate. CH mass showed a genetic correlation with leaf mass. Developmental plasticity in modular allocation allows Microstegium to maximize fitness when conditions are favorable (e.g., high light along forest edges) by continual maturation of CL caryopses on axillary racemes.     

Life history trade-offs in Amphibromus scabrivalvis (Poaceae): allocation to clonal growth,storage, and cleistogamous reproduction

Life history trade-offs among clonal growth, storage, and sexual reproduction were investigated in the perennial grass Amphibromus scabrivalvis in relation to soil nutrients. This species exhibits clonal growth by producing rhizomes and stores reserves in the form of basal corms; seeds are matured in cleistogamous spikelets on panicles enclosed within the leaf sheaths along each culm. Ten seed-derived genotypes (clones) were separated into 72 ramets and planted in the greenhouse. Control ramets received only water while the remainder received fertilizer applied every 2 wk. Twenty-four ramets were harvested per clone at 11, 20, and 26 wk. The dry mass of corms, rhizomes, roots, shoots, and seeds were recorded. Biomass partitioning to rhizomes provided a measure of carbon allocation to clonal growth, partitioning to corms provided a measure of allocation to storage, and partitioning to seeds provided a measure of allocation to sexual reproduction. Allocation to most organs was significantly influenced by clone identity; fertilizer significantly increased allocation to corms and seeds at 20 wk, but never affected rhizome allocation at any age. Corm allocation increased from 2% at 11 wk to 27% at 26 wk; rhizome allocation decreased from 10% at 11 wk to 3% at 26 wk. Significant negative relationships were detected for rhizome vs. seed and corm vs. rhizome allocation in fertilized clones at 20 wk. This suggests an age-dependent physiological life history trade-off between clonality and sexual reproduction and between clonality and storage. In contrast, a significant positive relationship was consistently noted for corm vs. seed allocation in fertilized and unfertilized clones at 20 and 26 wk. The absence of a trade-off between storage and sexual reproduction may indicate that these two processes are not necessarily mutually exclusive components of life history.     

Recovery from drought stress in Lolium perenne (Poaceae): are fungal endophytes detrimental?

Perennial ryegrass (Lolium perenne) is a cool-season, perennial species widely used for forage and turf. It is often infected by a clandestine, endophytic fungus (Neotyphodium lolii) that has the potential to affect host growth responses to abiotically stressful conditions. In some species, the grass-endophyte symbiosis is mutualistic, but the relationship is reported to be contingent on environmental conditions and host genotype in L. perenne. The objective of this research was to determine the potential effects of endophyte infection on recovery from severe drought stress in variable genotypes of a perennial ryegrass cultivar. Sixteen infected (+E) and 16 uninfected (-E) ramets were planted in the greenhouse for each of 10 ryegrass genotypes. Eight +E and eight -E plants per genotype were exposed to three sequential droughts where water was withheld for 11-14 d, resulting in <5% soil moisture; the others (control) were watered as needed. Response variables were tiller numbers 1 wk and 4 wk after drought, and leaf area and dry mass of shoots and roots 7 wk after drought. In both control and drought, -E plants had more tillers, and greater leaf area and total mass, than +E plants, suggesting a detrimental effect of endophytic fungi. Fungal hyphae survived the drought and were abundant in post-drought, +E plants. The effects of endophytes were specific for particular host genotypes, as exemplified by significant genotype × endophyte interactions. Root : shoot ratio and percent of mass allocated to tiller bases (a rough measure of resource storage) showed genotype × endophyte × drought interactions. There was plasticity for root : shoot ratio and genetic variation in the ability to restore root growth during recovery from drought. For 7 of 10 genotypes, -E plants showed an equal or greater allocation to tiller bases than +E plants following drought recovery, illustrating a cost to endophyte infection for some genotypes. The symbiotic relationship between L. perenne and its endophyte primarily benefits the fungus, not the host, under many environmental conditions.     

BIRD-DISPERSAL OF PHYTOLACCA AMERICANA L. AND THE INFLUENCE OF FRUIT REMOVAL ON SUBSEQUENT FRUIT DEVELOPMENT

Characteristics of fruits, seeds, and ripening of Phytolacca americana were studied in New Jersey, USA to assess their importance in the dispersal strategy of this species. Removal by birds was directly related to the percent of ripe fruits available on a raceme. Fruit removal from a raceme resulted in lower fruit pulp and seed weights for fruits developing on the same raceme, but these differences probably do not influence probability for dispersal by birds. We demonstrate that characteristics of the fruit pulp may decrease seed availability for the primary seed predator, Peromyscus leucopus, and increase the time fruits are available for dispersal. We speculate on the significance of the sequential fruiting in Phytolacca and the relationship to availability of dispersal agents and high-lipid fruits. Sequential ripening and protection from predation result in seed dispersal from late August to early December.