Why did heterospory evolve?

The primitive land plant life cycle featured the production of spores of unimodal size, a condition called homospory. The evolution of bimodal size distributions with small male spores and large female spores, known as heterospory, was an innovation that occurred repeatedly in the history of land plants. The importance of desiccation‐resistant spores for colonization of the land is well known, but the adaptive value of heterospory has never been well established. It was an addition to a sexual life cycle that already involved male and female gametes. Its role as a precursor to the evolution of seeds has received much attention, but this is an evolutionary consequence of heterospory that cannot explain the transition from homospory to heterospory (and the lack of evolutionary reversal from heterospory to homospory). Enforced outcrossing of gametophytes has often been mentioned in connection to heterospory, but we review the shortcomings of this argument as an explanation of the selective advantage of heterospory. Few alternative arguments concerning the selective forces favouring heterospory have been proposed, a paucity of attention that is surprising given the importance of this innovation in land plant evolution. In this review we highlight two ideas that may lead us to a better understanding of why heterospory evolved. First, models of optimal resource allocation – an approach that has been used for decades in evolutionary ecology to help understand parental investment and other life‐history patterns – suggest that an evolutionary increase in spore size could reach a threshold at which small spores yielding small, sperm‐producing gametophytes would return greater fitness per unit of resource investment than would large spores and bisexual gametophytes. With the advent of such microspores, megaspores would evolve under frequency‐dependent selection. This argument can account for the appearance of heterospory in the Devonian, when increasingly tall and complex vegetative communities presented competitive conditions that made large spore size advantageous. Second, heterospory is analogous in many ways to anisogamy. Indeed, heterospory is a kind of re‐invention of anisogamy within the context of a sporophyte‐dominant land plant life cycle. The evolution of anisogamy has been the subject of important theoretical and empirical investigation. Recent work in this area suggests that mate‐encounter dynamics set up selective forces that can drive the evolution of anisogamy. We suggest that similar dispersal and mating dynamics could have underlain spore size differentiation. The two approaches offer predictions that are consistent with currently available data but could be tested far more thoroughly. We hope to re‐establish attention on this neglected aspect of plant evolutionary biology and suggest some paths for empirical investigation.     

Effect of light on abscisic acid content in photosensitive Scots pine (Pinus sylvestris L.) seed

Dormancy-breaking treatment of the photosensitive Scots pine (Pinus sylvestris L.) seed by white light incubation or a 15-min exposure to red light decreased the abscisic acid content prior to radicle protrusion. Incubation in the dark or exposure to red light followed by a 5-min far-red light irradiation did not cause as great a decrease in abscisic acid content nor was the dormancy relieved. The ability of the far-red light to keep the ABA level high and to prevent germination gradually disappeared as the length of the dark period between the red and far-red treatments was increased to 24 h. ABA was quantified on a gas chromatograph with an electron capture detector.     

Deposition and longevity of oviposition-deterring pheromone in the cabbage seed weevil

Abstract The cabbage seed weevil ( Ceutorhynchus assimilis Payk.) lays eggs singly into pods of oilseed rape ( Brassica napus L.) through punctures bored with the mouthparts, preferring pods not recently used for oviposition. A simple new choice test has been used to test individual components of egg-laying behaviour for their effect on oviposition site selection. It is confirmed that an oviposition-deterring pheromone (ODP) is deposited during abdomen brushing of the pod which follows egg-laying. Neither pin punctures, weevil feeding punctures, oviposition punctures nor eggs had any deterrent effect. Pods walked on by female weevils were not avoided by those laying eggs. Observations suggest that the ODP is sensed by contact chemoreceptors on the antennae. The deterrent effect lasted only 1–2 h. The implications of these findings on the adaptive significance of the pheromone and its possible use in pest control are discussed.     

The endotegmen tuberculae: an account of little-known structures from the seed coat of the Hydrocharitoideae (Hydrocharitaceae) and Najas (Najadaceae)

SHAFFER-FEHRE, M., 1991. The endotegmen tuberculae: an account of little-known structures from the seed coat of the Hydrocharitoideae (Hydrocharitaceae) and of Najas (Najadaceae). The sub-cellular protuberances from the inner tegmen layer have been given various names in earlier literature. Here named 'endotegmen tuberculae', they vary in shape, size and density of distribution in the endotegmen of different taxa. These characteristics make them useful for taxonomic research at both genus and species level. Their shape is a major consideration in transferring Hydrocharis dubia to the genus Limnobium. A further structure, 'exotegmen tuberculae', has been observed for the first time. This paper gives a survey of these structures and illustrates their taxonomic importance and its implications.     

Spatial metrics for detecting ecosystem degradation in the ridge-slough patterned landscape

Indicators of landscape condition should be selected based on their sensitivity to environmental changes and their capacity to provide early warning detection of those changes. We assessed the performance of a suite of spatial-pattern metrics selected to quantify the condition of the ridge-slough landscape in the Everglades (South Florida, USA). Spatial pattern metrics (n = 14) that describe landscape composition, geometry and hydrologic connectivity were enumerated from vegetation maps of twenty-five 2 × 2 km primary sampling units (PSUs) that span a gradient of hydrologic and ecological condition across the greater Everglades ecosystem. Metrics were assessed in comparison with field measurements from each PSU of landscape condition obtained from regional surveys of soil elevation, which have previously been shown to capture dramatic differences between conserved and degraded locations. Elevation-based measures of landscape condition included soil elevation bi-modality (BI_SE), a binary measure of landscape condition, and also the standard deviation of soil elevation (SD_SE), a continuous measure of condition. Metric performance was assessed based on the strength (sensitivity) and shape (leading vs. lagging) of the relationship between spatial pattern metrics and these elevation-based measures. We observed significant logistic regression slopes with BI_SE for only 4 metrics (slough width, ridge density, directional connectivity index – DCI, and least flow cost – LFC). More significant relationships (n = 8 metrics) were observed with SD_SE, with the strongest associations for slough density, mean ridge width, and the average length of straight flow, as well as for a suite of hydrologic connectivity metrics (DCI, LFC and landscape discharge competence – LDC). Leading vs. lagging performance, inferred from the curvature of the association obtained from the exponent of fitted power functions, suggest that only DCI was a leading metric of the loss of soil elevation variation; most metrics were indeterminate, though some were clearly lagging. Our findings support the contention that soil elevation changes from altered peat accretion dynamics precede changes in landscape pattern, and offer insights that will enable efficient monitoring of the ridge-slough landscape as part of the ongoing Everglades restoration effort.     

Flow cytometric,chromosomal and morphometric analyses challenge current taxonomic concepts in the Portulaca oleracea complex (Portulacaeae,Caryophyllales)

Portulaca oleracea is a noxious annual weed of worldwide distribution in temperate to tropical climates. Its taxonomy has been treated in contradictory ways in the past. Various microspecies have been described, lumped into a single species by other authors. We re‐examined the importance of seed size and ploidy variation, previously applied as the most important taxonomic characters, for systematic classification based on accessions from Europe, Asia, Africa and South America using flow cytometry, chromosome counting and morphometry. Sixteen microspecies and six transitional forms, covering the ploidy and seed character variation, proposed for the complex, were studied from 178 populations. Portulaca grandiflora was included as a reference species from outside the complex. DNA hyper‐pentaploidy or hexaploidy were inferred for the majority of accessions which exhibited the full range of seed size. It is recommended that the only species of lower ploidy (either diploid based on x = 12 or tetraploid based on x = 12) encountered, P. nicaraguensis, should be separated from the P. oleracea complex as it deviates in base chromosome number and monoploid genome size. The frequency distribution of seed size was continuous and unimodal within the wild taxa of the complex and in pairs of taxa defined by testa sculpture. Seed size of DNA hexaploids was slightly negatively correlated with sample/standard fluorescence intensities. Our results conflict with the current microspecies concept. Possible reasons underlying the discrepancy are discussed and strategies for future systematic research are suggested. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 144–156.     

Stress integrated tests and cytological analyses reveal Brassica villosa subsp. drepanensis seed quality decrease upon long-term storage

Under stress integrated germination test (SIGT), seeds undergo osmo-saline stresses, which enable to detect differences in vigour of long-term stored seeds with high germination percentage (G%). The quality of Brassica villosa subsp. drepanensis seeds stored in a genebank (at ? 20°C for 16 years) was compared with seeds at harvest by standard germination tests (GT), SIGT and cytogenetic analysis. No differences were detected in G% and mean germination time under GT. Conversely, SIGT performed with NaCl ? 0.9 MPa osmotic potential did not influence G% at harvest but reduced that of stored seeds, SIGT at ? 1.4 MPa reduced G% of both. Cytogenetic analysis showed reduction of mitotic index, appearance of chromosomal aberrations and smaller nucleoli in stored seeds compared with harvest seeds germinated in water. SIGT at ? 0.9 MPa had no effect on mitotic index, but increased chromosome aberrations and nucleoli number. SIGT at ? 1.4 MPa inhibited G% of harvest and stored seeds, reduced mitoses in harvest and completely prevented it in stored seeds. The results indicate that GT does not faithfully reflect the quality of stored seeds, with misinterpretation of their vigour, whereas SIGT and cytogenetical parameters are sensitive, reliable and inexpensive methods for early prediction of genetic erosion in germplasm banks.     

Wetland drying indirectly influences plant community and seed bank diversity through soil pH

High altitude wetlands on the Tibetan Plateau have been shrinking due to anthropogenic disturbances and global climate change. However, the few studies that have been conducted on wetlands are inconclusive about the effect of soil moisture on seed banks and potential of seed banks in wetlands with different levels of soil moisture for regeneration of dried wetlands. We investigated seed banks and plant communities along a soil moisture gradient. A structural equation model was used to analyze the direct and indirect effects of soil moisture on seed banks, as well as the relationship between plant communities and seed banks. Although soil moisture had no direct effects on seed bank richness and density and indirect effects on seed banks through plant community, it had indirect effects on the seed bank through soil pH. Soil moisture also did not have direct effects on plant community richness, but it had indirect effects through soil pH. Plant community composition changed with soil moisture, but aboveground plant abundance and seed banks composition did not change. Low similarity exists between plant community and seed banks for all wetlands, and similarity decreased along the moisture gradient. The key factor determining plant community diversity was soil pH, while seed bank diversity was mainly affected by soil pH and plant community diversity with wetland drying. Although potential for regenerating the plant community from the seed bank decreased with an increase in soil moisture, drained wetlands still have enough residual seeds for successful restoration of species-rich alpine meadows.     

Transferring sediment containing intact seed banks: a method for studying plant community ecology

The experimental study of the ecology of natural plant communities is necessary to demonstrate the effects of environmental factors on plant growth. It allows one to make predictions of the likely effects of environmental changes on plant communities. Manipulating an entire ecosystem is complicated, and is rarely attempted. An alternative method is applied here for the submerged plant communities of the seasonally-flooded oligohaline marshes of the Camargue. It consists of collecting undisturbed sediment samples with their contained intact seed bank, and submitting them to different salinities. The study of total biomass and of the biomass of the more frequent species was carried out using parametric tests. The precision of the results depended on the frequency of the species in the replicates. When the frequency was low, non-parametric statistics were necessary. The method seems to be particularly suitable for the study of communities of annual species in aquatic environments.

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Résumé L'étude expérimentale de l'écologie des communautés naturelles de plantes est nécessaire pour démontrer les relations de cause à effet entre les facteurs environnementaux et l'abondance des plantes. Elle l'est également pour faire des prédictions réalistes sur les effets de modifications envisagées de l'environnement sur des communautés de plantes. La manipulation de l'écosystème complet est complexe, ce qui rend son utilisation rare. Une méthode alternative a été utilisée pour l'étude de la composition spécifique des communautés de plantes submergées des marais temporaires de Camargue. Elle consiste à prélever des échantillons non perturbés de sédiment, avec leur stock de semences intact, et à les soumettre à des conditions de salinité. L'étude de la biomasse totale et de la biomasse des espèces les plus fréquentes est faite au moyen de test paramétriques. La précision attendue des résultats dépend de la fréquence de l'espèce dans les réplicats. Lorsque la fréquence n'est pas élevée l'emploi de statistiques non paramétriques est indispensable. La méthode se révéle particulièrement bien adaptée à l'étude des communautés d'annuelles en milieu aquatique.