Evolution and ecology of seed internal morphology in relation to germination characteristics in Amaranthaceae.

BACKGROUND AND AIMS: Internal seed morphological traits such as embryo characteristics and nutritive tissue can vary considerably within a plant lineage. These traits play a prominent role in germination processes and the success of seedling establishment, and are therefore under high selective pressure, especially in environments hostile to seedlings, such as arid, saline or highly dynamic habitats. We investigated the relationships of seed internal morphology and germination characteristics of 84 species of Amaranthaceae s.l., a family with numerous lineages that have adapted to stressful growing conditions. METHODS: We used seed cross-sections to assess embryo type and the ratios of embryo to seed surface and radicle to cotyledon length. Furthermore, seed mass, mean time to germination, habitat preferences and further plant traits such as C3 or C4 photosynthesis and life form were compiled for each species. Data were analysed using phylogenetic comparative methods. KEY RESULTS: We found embryo type (λ = 1), log seed mass (λ = 0.86) and the ratio of embryo to seed size (λ = 0.78) to be evolutionarily stable, with an annular embryo as ancestral in the family. Linked to shifts to the three derived embryos types (spiral, horseshoe-shaped and curved) is an increase in the ratio of root to cotyledon length and a reduction of nutritive tissue. We observed stabilizing selection towards seeds with relatively large embryos with longer radicles and less nutritive tissue that are able to germinate faster, especially in lineages with C4 photosynthesis and/or salt tolerance. CONCLUSIONS: We conclude that the evolutionary shift of nutrient storage from perisperm to embryo provides an ecological advantage in extreme environments, because it enables faster germination and seedling establishment. Furthermore, the evolutionary shift towards a higher ratio of root to cotyledon length especially in small-seeded Amaranthaceae growing in saline habitats can provide an ecological advantage for fast seedling establishment.

C4-like photosynthesis and the effects of leaf senescence on C4-like physiology in Sesuvium sesuvioides (Aizoaceae).

Sesuvium sesuvioides (Sesuvioideae, Aizoaceae) is a perennial, salt-tolerant herb distributed in flats, depressions, or disturbed habitats of southern Africa and the Cape Verdes. Based on carbon isotope values, it is considered a C4 species, despite a relatively high ratio of mesophyll to bundle sheath cells (2.7:1) in the portulacelloid leaf anatomy. Using leaf anatomy, immunocytochemistry, gas exchange measurements, and enzyme activity assays, we sought to identify the biochemical subtype of C4 photosynthesis used by S. sesuvioides and to explore the anatomical, physiological, and biochemical traits of young, mature, and senescing leaves, with the aim to elucidate the plasticity and possible limitations of the photosynthetic efficiency in this species. Assays indicated that S. sesuvioides employs the NADP-malic enzyme as the major decarboxylating enzyme. The activity of C4 enzymes, however, declined as leaves aged, and the proportion of water storage tissue increased while air space decreased. These changes suggest a functional shift from photosynthesis to water storage in older leaves. Interestingly, S. sesuvioides demonstrated CO2 compensation points ranging between C4 and C3-C4 intermediate values, and immunocytochemistry revealed labeling of the Rubisco large subunit in mesophyll cells. We hypothesize that S. sesuvioides represents a young C4 lineage with C4-like photosynthesis in which C3 and C4 cycles are running simultaneously in the mesophyll.

C3 -C4 intermediates may be of hybrid origin - a reminder.

The currently favoured model of the evolution of C4 photosynthesis relies heavily on the interpretation of the broad phenotypic range of naturally growing C3 -C4 intermediates as proxies for evolutionary intermediate steps. On the other hand, C3 -C4 intermediates had earlier been interpreted as hybrids or hybrid derivates. By first comparing experimentally generated with naturally growing C3 -C4 intermediates, and second summarising either direct or circumstantial evidence for hybridisation in lineages comprising C3 , C4 and C3 -C4 intermediates, we conclude that a possible hybrid origin of C3 -C4 intermediates deserves careful examination. While we acknowledge that the current model of C4 photosynthesis evolution is clearly the best available, C3 -C4 intermediates of hybrid origin, if existing, should not be used for further analysis of this model. However, experimental C3  × C4 hybrids potentially are excellent systems to analyse the genetic differences between C3 and C4 species and, also using segregating progeny, to study the relationship between individual photosynthetic traits and environmental factors.