Limited seed dispersal and microspatial population structure of an agamospermous grass of West African savannahs, Hyparrhenia diplandra (Poaceae)

We studied the microspatial population structure of the perennial tussock grass, Hyparrhenia diplandra (Poaceae), a facultative agamospermous species of West African savannahs. The microspatial population structure of H. diplandra was investigated by choosing two 100-m(2); quadrats at random from which all individuals were mapped. The genotype of every individual was determined using two highly polymorphic microsatellite loci. A chloroplast locus was also used to investigate the role of seed dispersal on the genetic structure of populations. The genetic diversity index (0.85) was high for a clonal species. Significant genetic differentiation over short distances was detected by F statistics, and spatial autocorrelation analyses within both quadrats showed significant isolation-by-distance patterns, both with the cytoplasmic locus and the nuclear loci. Some clones formed large patches (up to 5 m in diameter) whereas others were more scattered. However, the genetic differentiation between quadrats was much higher when studied with the cytoplasmic locus than with the nuclear loci, indicating that gene flow via pollen, but not seeds, may frequently occur between quadrats. The maintenance of genetic diversity in this facultative agamospermous species most likely results from several factors, such as low seed dispersal ability, nonnegligible gene flow through pollen, and selective pressures induced by regularly occurring fires in this ecosystem.     

Relationships between root density of the African grass Hyparrhenia diplandra and nitrification at the decimetric scale: an inhibition-stimulation balance hypothesis

Previous studies have shown that Lamto savannah exhibits two different types of nitrogen cycle with high and low nitrification sites and suggested that the perennial grass Hyparrhenia diplandra is responsible for this duality at a subpopulation level, with one ecotype being thought to be able to inhibit nitrification. The present work aimed to investigate the relationships between nitrification and the roots of H. diplandra at two scales. (i) Site-scale experiments gave new insight into the hypothesized control of nitrification by H. diplandra tussocks: the two ecotypes exhibited opposite influences, inhibition in a low nitrification site (A) and stimulation in a high nitrification site (B). (ii) Decimetric-scale experiments demonstrated close negative or positive relationships (in sites A or B, respectively) between the roots and nitrification (in the 0-10 cm soil layer), showing an unexpectedly high sensitivity of the nitrification process to root density. In both soils, the correlation between the roots and nitrification decreased with depth and practically disappeared in the 20-30 cm soil layer (where the nitrification potential was found to be very low). Therefore, the impact of H. diplandra on nitrification may be viewed as an inhibition-stimulation balance.     

Reduced recombination in gynodioecious populations of a facultative apomictic orchid

Many plants combine sexual reproduction with some form of asexual reproduction to different degrees, and lower genetic diversity is expected with asexuality. Moreover, the ratios of sexual morphs in species with gender dimorphism are expected to vary in proportion to the reproductive success of the sexual process. Hence, sex ratios can directly influence the genetic structure and diversity of a population. We investigated genotypic diversity in 23 populations of a facultative, apomictic gynodioecious orchid, Satyrium ciliatum, to examine the effect on genotypic diversity of variation in the frequency of females and in the amount of sexual reproduction. The study involved one pure female, seven gynodioecious (both females and hermaphrodites present) and 15 hermaphroditic populations. Pollinia receipt was higher in hermaphroditic than in gynodioecious populations. Analyses of variation in ISSRs demonstrated that genotypic diversity was high in all populations and was not significantly different between hermaphroditic and gynodioecious populations. We used character compatibility analysis to determine the extent to which recombination by sexual reproduction contributed to genotypic diversity. The results indicate that the contribution of recombination to genotypic diversity is higher in hermaphroditic than in gynodioecious populations, consistent with the finding that hermaphroditic populations received higher amounts of pollinia. Our finding of reduced recombination in gynodioecious populations suggests that maintenance of sex in hermaphrodites plays an important role in generating genotypic diversity in this apomictic orchid.     

Genetic fingerprinting for determining the mode of reproduction in Paspalum notatum, a subtropical apomictic forage grass

 Paspalum is an important genus of the family Gramineae that includes several valuable forage grasses. Many of the species are polyploid and either obligate or facultative apomicts. Cyto-embryological observations of several tetraploid genotypes of P. notatum were performed to determine their mode of reproduction. Afterwards, selfed progenies of the genotypes F131, Q3664 and Q4117 were analysed using RFLP and RAPD genetic fingerprints to identify maternal and non-maternal (aberrant) plants, and to establish the degree of apomictic reproduction. Five maize clones and six primers were used for detecting genetic deviations from the maternal profile. Maize clones umc379, umc384 and umc318 and primers OPG10 and OPI4 were the most informative for discriminating between maternal and aberrant individuals within the progenies of F131 and Q3664. The combined results of three RFLP clones or 4–6 RAPD primers were necessary to ascertain the mode of reproduction in plants F131 and Q3664. The results obtained with the RFLP and RAPD markers were in agreement with the cyto-embryological studies in ascertaining the mode and degree of apomictic reproduction. Plant F131 showed a completely sexual reproductive behaviour, Q3664 an elevated expression of sexuality, while Q4117 was highly apomictic. A fingerprint analysis of an outcrossing population, aimed at the identification of hybrid plants, was also performed. Maize clones um318 and umc379 and primers OPC2 and OPC9 were used. The presence of specific bands belonging to the male parent permitted a rapid and easy detection of hybrids. The methodology described here can be applied both for the characterisation of P. notatum populations and to identify hybrid progenies in Paspalum breeding programs. Received: 5 March 1997 / Accepted: 13 May 1997     

Determining genetic origins of aberrant progeny from facultative apomictic Kentucky bluegrass using a combination of flow cytometry and silver-stained RAPD markers

Seeded plants that reproduce through facultative apomixis produce two types of progeny: (1) apomictic progeny genetically identical to the maternal genotype, and (2) aberrant progeny genetically different from the maternal genotype. Aberrant progeny have at least nine different genetic origins depending on gametic ploidy level and whether fertilization was self, cross, or absent. Multiple genetic origins of aberrant progeny complicate the results of basic and applied genetic studies. Determining the genetic origin of progeny plants using traditional techniques, such as cytology, embryology, and segregational studies, is technically difficult in Kentucky bluegrass. We have found that two relatively new techniques, flow cytometry and silver-stained RAPD (ssRAPD) markers, are powerful tools for rapidly determining the genetic origins of aberrant Kentucky bluegrass progeny. Our application of these techniques demonstrate that (1) flow cytometry accurately distinguishes progeny ploidy levels, and (2) ssRAPD markers distinguish progeny resulting from cross-fertilization. Therefore, a combination of flow cytometry and ss-RAPD data would be useful for most genetic studies of aberrant individuals. Moreover, ssRAPD s were found to be of value for measuring the loss of genetic markers from polyhaploids and quantifying the inheritance of parental genomes in polydiploid B_n (n+n) and polytriploid B_III (2n+n) hybrids. Quantifying shared ss-RAPD markers may also be useful for determining genetic relatedness between varieties and germplasm sources.     

Modelling the recovery of an annual savanna grass following a fire-induced crash

Abstract The native annual Sorghum populations of the Australian wet-dry tropics are highly resilient to dry season fires. During the early wet season, however, fires that occur after the new grass population has emerged can cause catastrophic population crashes. We examined savanna plots that had been burnt in this way, and compared them with adjacent unburnt plots. We found that Sorghum densities in the burnt plots were lower on average by a factor of 10, but that some fires had reduced the density only to one-third of the unburnt plots. It is not clear whether these differences relate directly to site or seasonal factors, or to differences in the way the burning was carried out. Other vegetation components responded to the fires differently: forbs (dicotyledonous herbs) increased in cover, while perennial grasses, woody plants, and overall species richness, were not significantly affected. The amount of leaf litter declined. A population model for Sorghum based on the demography of unburnt populations predicted that they should recover from a wet season burn, taking 7–16 years to return to normal densities. However, the actual field populations did not seem to be recovering, suggesting that wet season fires not only lower densities, but may also fundamentally change population processes in these annual grasses.     

Contrasting impacts of grass species on nitrogen cycling in a grazed Sudanian savanna

We investigated the impact of perennial and annuals grass species on nitrogen cycling in a Sudanian savanna of Burkina Faso. We also analysed how the local context in terms of grazing and soil properties modifies these impacts. We selected four plots differing both by the intensity of grazing by cattle and soil depth, and used soil and grass biomass ¹⁵N as integrative indicators of N cycle. If perennials are able to foster a more efficient nitrogen cycling there should be lower ¹⁵N abundances in their biomass and soil. If soil depth and cattle pressure significantly modify nitrogen fluxes, soil depth and cattle pressure should influence ¹⁵N signatures. Our results suggest that perennial grasses are more conservative for nitrogen (inhibition of nitrification, less leaching via a perennial root system, slower cycling). The increase in leaf δ¹⁵N with N concentration is steeper in Loudetia togoensis than in the three other grasses. No significant difference was found between the ¹⁵N signatures of the four plots. Our results on ¹⁵N signatures and the fact that perennial grasses are much more abundant in the plots that are less grazed and have deeper soils, confirm that the switch from perennial to annual grasses is linked to a degradation in soil fertility and pasture quality. This suggests that ¹⁵N signatures can be used as indicators of fertility.     

Architecture,seasonal growth and interference in three grass species with different flowering phenologies in a tropical savanna

In a field experiment we studied the relationships between architecture, seasonal aerial biomass growth and interference from neighbours in three savanna grass species differing in time of flowering: a precocious species (Elyonurus adustus), an early species (Leptocoryphium lanatum) and a late species (Andropogon semiberbis). To detect the neighbour interference upon architecture and seasonal regrowth, we measured the leaf interception of a plane at different heights, when the species grew alone, in pairs and in groups of three. Although the three species differed widely in the spatial and temporal patterns of occupation of above-ground space, important levels of interference among neighbouring plants were detected. The species differ in: a) the pattern of space occupation when growing alone; b) the magnitude of the interference effect by the companion species; c) the change in pattern as a consequence of interference; d) their reaction to fire.The interference is not symmetric and it is not related to phenological similarities based on flowering season.L. lanatum was both the most impaired and the least impairing of the three species and the opposite is valid forA. semiberbis. The basal species (E. adustus andL. lanatum) showed a higher potential to produce leaf surface during the growth season than the tall species (A. semiberbis), but the stronger interference from the latter tended to equate their growth when the species grew in mixtures.In all three species interference led to a reduction of the growing period. Based on the analysis of growth at the various heights and the architectural peculiarities of the species we concluded that neighbour interference is probably reducing both tillering and leafing in the basal species but only culm elongation and leafing in the erect species.The species also differed in their reaction to fire.A. semiberbis andE. adustus showed a pulse of regrowth after the passing of fire which is missing inL. lanatum. In all cases the neighbour interference affected both the intensity of growth and the length of the growing period, but it did not affect either the height of the plants nor the reproductive phenology of these three grass species.     

Herbivore and nutrient control of lawn and bunch grass distributions in a southern African savanna

In African savannas, bottom-up soil nutritional factors and top-down herbivory have both been suggested to control the distribution patterns of bunch and lawn grasses. We tested the separate and combined roles of these factors on grassland distribution in a South African savanna by focusing on three main objectives, namely (1) are grazing patches and lawns restricted to specific soils or sites, (2) does herbivory alter rates of nutrient cycling which facilitates lawns and (3) are there any differences in foliage quality between lawn and bunch grass-dominated sites that might influence animals in choosing to feed on lawns? We set up ten sites along a grazing gradient in the Hluhluwe-iMfolozi Park in KwaZulu-Natal, South Africa. At these sites, we measured total soil nutrients, soil nutrient turnover rates and grass foliage nutrient concentrations. We found that in the Hluhluwe-iMfolozi Park the spatial distribution of lawn and bunch grassland types appears to be an animal-driven phenomenon and not dependent on specific soil properties. The short-structured and distinct species found in lawns do not appear to be restricted to nutrient-enriched patches. However, the grasses of these lawns had significantly higher nutrient concentrations, in their foliage, which might explain the high attraction to these patches by herbivores. We also did not find any animal-induced stimulation of nutrient cycling rates that are often associated with lawn grass species.     

Belowground competitive suppression of seedling growth by grass in an African savanna

Coexistence of N₂-fixing legumes and non-legume trees with grasses in African savannas results in intense competition between these life-forms. We hypothesised that belowground competition might induce different nutritional constraints in N₂- versus non-N₂-fixing species. A field (Hluhluwe-imFolozi nature reserve, South Africa) competition experiment with two N₂-fixing legume species (Acacia burkei and Acacia karroo) and two non-N₂-fixing species (Schotia brachypetala and Spirostachys africana) both with clipped grass and without grass was established. Plants were supplied with no fertilizer, or generous amounts of fertilizer (200?kg?N?ha^?1, 100?kg P₂O₅ ha^?1, 7.1?kg K₂O ha^?1) supplied as either 28?C10 (N?CK), P or a combination of these fertilizers (NPK). Regularly clipped grass suppressed growth (by more than 90?%) of both N₂- and non-N₂-fixing seedlings equally. Biomass accumulation of seedlings grown with grass and the grasses themselves responded positively to NK and/or NPK, but not P, although P-fertilization did have effects on foliar [N] and ??¹⁵N values of trees and grasses showing that plants accessed the fertilizer. Tree ??¹⁵N values and foliar [N] were also modified by NPK, demonstrating access to fertilizer. However, the ameliorative effects of NPK on grass competition-induced biomass suppression were only partial. This may be due to ??non-resource competition?? (i.e. root gaps) imposed by dense grass roots. The fact that nutrients were able to partially ameliorate the effects of grass competition, however, indicates that such ??non-resource competition?? may be partially overcome by even more generous supply of fertilizers.     

Developmental expression of ASG- 1 during gametogenesis in apomictic guinea grass (Panicum maximum)

We have used Western blue-visualized in situ-hybridization (ISH) to monitor the expression of apomixis-specific gene-1 (ASG-1, GenBank accession number AB000809) during gametogenesis in obligate-sexual and facultative-apomictic (aposporic) genotypes of guinea grass (Panicum maximum). The in situ-analysis revealed that ASG-1 is not expressed in the ovule during early floral development in both, the facultative apomicts (A1 stage) and the obligate sexuals (S1 stage). With the appearance of the aposporous initial cell(s) in the ovule of the apomictic type (A2-1 stage), ASG-1 expression is strong and specific to this apomixis-specific cell. ASG-1 expression continued through different stages of aposporous embryo sac development (A2-2 stage), indicating that the gene may play a role in this developmental process. Regular embryo sacs in sexual types did not show hybridization signals (S2 stage). However, strong ASG-1 expression was detected in immature pollen grains and young embryos in both reproductive types, suggesting that ASG-1 may be an allele derived from the obligate-sexual wild type. Expression in pollen grains faded with maturation. In a heterologous system, using Paspalum notatum, a facultative-aposporic tropical grass (bahia grass), identical results were obtained. The results are discussed in view of the fact, that ASG-1 shows some homology to genes known to be seed- or embryo-specific or involved in processes related to cell growth.     

Seasonal selection of soil types and grass swards by roan antelope in a South African savanna

Roan antelope are distributed mainly in regions characterized by infertile soils, offering food of low quality. We hypothesized that roan may select localities with higher soil nutrient levels and/or grass swards with more favourable properties in terms of food abundance or quality than generally available in those regions. Roan antelope were observed in a savanna region in South Africa where soils of widely varying nutrient status occurred. Roan favoured open grassland over wooded savanna areas. During the wet season, roan preferred sites with felsite‐derived soil of intermediate soil nutrient status. Grasslands growing on nutrient enriched alluvial soils were preferred outside of the early wet season, although most of the favourable sward characteristics were present in other landscape units. Food quantity, rather than quality appeared to attract roan to foraging sites in the late wet and early dry seasons. Food quality appeared more important in the early wet and late dry seasons. The higher degree of clustering of leafy material within foraging swards seemed to be an additional discriminating factor. The factors governing the selection of foraging sites by roan did not seem notably different from those influencing other species of grazing ruminant, but roan nevertheless seemed tolerant of stemmy grasslands growing on nutrient richer substrates.     

Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna

Most savanna water balance models assume water partitioning between grasses and shrubs in a two-layer hypothesis, but this hypothesis has not been tested for humid savanna environments. Spatial partitioning of soil water between grasses and shrubs was investigated in a West African humid savanna by comparing the isotopic composition (oxygen-18 and deuterium) of soil water and plant stem water during rainy and dry conditions. Both grass and shrub species acquire most of their water from the top soil layer during both rainy and dry periods. A shift of water uptake pattern towards deeper horizons was observed only at the end of the dry season after shrub defoliation. The mean depth of water uptake, as determined by the isotopic signature of stem water, was consistent with grass and shrub root profiles and with changes in soil water content profiles as surveyed by a neutron probe. This provides evidence for potentially strong competition between shrubs and grasses for soil water in these humid savannas. Limited nutrient availability may explain these competitive interactions. These results enhance our understanding of shrub-grass interactions, and will contribute to models of ecosystem functioning in humid savannas.     

High-frequency direct plant regeneration via multiple shoot induction in the apomictic forage grass Cenchrus ciliaris L.

Genetic improvement of the apomictic forage grass species Cenchrus ciliaris L. based on conventional breeding methods is difficult and time-consuming. However, in vitro genetic manipulation of such species would provide a promising approach. A rapid and high-frequency in vitro plant regeneration protocol is essential for successful application of transgenic technology. This study reports on such a rapid, high-frequency and genotype-independent plant regeneration protocol for C. ciliaris L. Using the multiple shoot induction approach, up to 20 shoots per explant could be induced from shoot tips cultured on MS (Murashige and Skoog) medium when supplemented with 3.0 mg L⁻¹ TDZ. Two cultivars (IGFRI-3108 and IGFRI-727) and three exotic germplasm accessions (EC-397670, EC397496, and EC397336) showed equivalent responses to the protocol. Shoot tips from 4-d-old in vitro grown seedlings were used as explants for multiple shoot induction. Regenerated shoots were cultured on MS medium supplemented with gibberellic acid (2.0 mg L⁻¹) for shoot elongation. The regenerated shoots were rooted on MS medium supplemented with indole-3-acetic acid (3.0 mg L⁻¹). When transferred to soil in pots, hardened plants displayed up to 85% survival under field conditions.     

Dynamic spatial partitioning and coexistence among tall grass grazers in an African savanna

Competitive relationships among mobile animals may be expressed through dynamically changing spatial relationships over different time frames. Less common species that are apparently inferior competitors may be able to coexist with more abundant species by concentrating in regions of the landscape little utilized by the former at spatio‐temporal scales from annual or seasonal ranges to the specific foraging localities exploited at different stages of the annual cycle. Spatial relationships may be influenced further by dependencies on other resources, predation risks and facilitatory interactions under certain conditions. Our study aimed to determine whether competition with more abundant zebra and buffalo restricted the abundance of sable antelope in a region where these three tall‐grass grazers overlapped in their herd distributions. We tracked the simultaneous movements of animals representing herds of these species over two dry seasons and one wet season using GPS‐GSM collars, and estimated seasonal or monthly range extents and their overlap. We also compared daily separation distances between these animals against the null pattern expected if their movements had been independent, and assessed how prior grazing by buffalo influenced the subsequent use of these localities by sable. The range of the sable herd was mostly separated from the seasonal range of the buffalo herd during the late dry season of 2006 and throughout the dry season of 2007. Seasonal home ranges of zebra herds overlapped partially with the range of the sable herd during most of the year. Even during times when their ranges overlapped, sable were rarely recorded within <1 km of the buffalo herd. Prior grazing by buffalo beyond a threshold level inhibited later use of these localities by sable, but the sable were nevertheless able to exploit places that were little utilized by buffalo at that time. Sable were less able to evade overlap with the small, mobile zebra herds, and hence more vulnerable to competitive exclusion by zebra than by buffalo. Our findings demonstrate how less abundant species can restrict competition from more abundant competitors through dynamic spatial partitioning in regions where their home ranges overlap.     

Mode of reproduction is detected by Parth1 and Sex1 SCAR markers in a wide range of facultative apomictic Kentucky bluegrass varieties

Gametophytic apomixis in Kentucky bluegrass (Poa pratensis L.) involves the parthenogenetic development of unreduced eggs from aposporic embryo sacs. Marker-assisted selection for the mode of reproduction in P. pratensis would avoid costly and time-consuming phenotypic progeny tests. We developed and tested two SCAR primer pairs that are associated with the mode of reproduction in P. pratensis. The SCAR primers identified the apomictic and sexual genotypes among progenies of sexual x apomictic crosses with very low bias. Furthermore, when tested on a wide range of Italian and exotic P. pratensis germplasm, they were able to unequivocally distinguish sexual from apomictic genotypes. This system should, therefore, allow new selection models to be set up in this species.