Evidence of combined flower thermal and drought vulnerabilities portends reproductive failure under hotter-drought conditions.

Despite the abundant evidence of impairments to plant performance and survival under hotter-drought conditions, little is known about the vulnerability of reproductive organs to climate extremes. Here, by conducting a comparative analysis between flowers and leaves, we investigated how variations in key morphophysiological traits related to carbon and water economics can explain the differential vulnerabilities to heat and drought among these functionally diverse organs. Due to their lower construction costs, despite having a higher water storage capacity, flowers were more prone to turgor loss (higher turgor loss point; ΨTLP) than leaves, thus evidencing a trade-off between carbon investment and drought tolerance in reproductive organs. Importantly, the higher ΨTLP of flowers also resulted in narrow turgor safety margins (TSM). Moreover, compared to leaves, the cuticle of flowers had an overall higher thermal vulnerability, which also resulted in low leakage safety margins (LSM). As a result, the combination of low TSMs and LSMs may have negative impacts on reproduction success since they strongly influenced the time to turgor loss under simulated hotter-drought conditions. Overall, our results improve the knowledge of unexplored aspects of flower structure and function and highlight likely threats to successful plant reproduction in a warmer and drier world.

Anemochorous and zoochorous seeds of trees from the Brazilian savannas differ in fatty acid content and composition.

Fatty acids (FAs) stored as triacylglycerols (TAGs) are an important source of carbon and energy for germination and seedling development, particularly for plants with small wind-dispersed seeds, allowing greater efficiency in storing both energy and carbon. These plants should be under strong selection to produce seeds rich in FAs and with large amounts of saturated FAs. Their closely packed single-chain configuration allows greater packing, more carbon and energy per unit mass, and are less costly to produce. Efficient carbon storage would be less crucial for zoochorous species, which can reach much larger seed sizes (mass). We analysed the transesterified FA profile from seeds of 22 anemochorous and zoochorous tree species from the Cerrado savannas of Central Brazil. We tested if seed FA content covaried with seed mass and if anemochorous and zoochorous seeds differed in FA contents and distribution. Fatty acids were an important seed source of carbon and energy for most species. Fifteen different FAs were identified. Oleic, linoleic and linolenic tended to be the predominant unsaturated FAs. Oleic acid corresponded to more than 60 % of the total transesterified FAs in seeds of Kielmeyera coriacea, Qualea dichotoma and Triplaris americana. Linoleic acid corresponded to more than 50 % of total FA in Dalbergia miscolobium, Parkia platycephala and Ferdinandusa elliptica while linolenic acid was the dominant component in Inga cylindrica. Across species, palmitic and stearic were the dominant saturated FAs. The only exception was lauric acid (68 % of total FA) in seeds of Qualea grandiflora. On a log10 scale, as the seed increased in mass, accumulation of FAs tends to proceed at a faster rate in anemochorous species than in zoochorous species. They also became increasingly richer in saturated FAs. Zoochorous species had seed TAGs with higher proportion of polyunsaturated FAs.

A Structure Shaped by Fire, but Also Water: Ecological Consequences of the Variability in Bark Properties Across 31 Species From the Brazilian Cerrado.

Bark is a structure involved in multiple physiological functions, but which has been traditionally associated with protection against fire. Thus, little is known about how the morpho-anatomical variations of this structure are related to different ecological pressures, especially in tropical savanna species, which are commonly subjected to frequent fire and drought events. Here we evaluated how the structural and functional variations of bark are related to the processes of resilience and resistance to fire, as well as transport and storage of water in 31 native species from the Brazilian Cerrado. Because of their thick bark, none of the trees analyzed were top-killed after a severe fire event. The structural and functional variations of the bark were also associated with water storage and transport, functions related to properties of the inner bark. In fact, species with a thicker and less dense inner bark were the ones that had the highest water contents in the wood, bark, and leaves. Lower bark density was also related to higher stem hydraulic conductivity, carbon assimilation, and growth. Overall, we provide strong evidence that in addition to protection from fire, the relative investment in bark also reflects different strategies of water use and conservation among many Cerrado tree species.

Flood tolerance in two tree species that inhabit both the Amazonian floodplain and the dry Cerrado savanna of Brazil.

Comparing plants of the same species thriving in flooded and non-flooded ecosystems helps to clarify the interplay between natural selection, phenotypic plasticity and stress adaptation. We focussed on responses of seeds and seedlings of Genipa americana and Guazuma ulmifolia to substrate waterlogging or total submergence. Both species are commonly found in floodplain forests of Central Amazonia and in seasonally dry savannas of Central Brazil (Cerrado). Although seeds of Amazonian and Cerrado G. americana were similar in size, the germination percentage of Cerrado seeds was decreased by submergence (3 cm water) and increased in Amazonian seeds. The seeds of Amazonian G. ulmifolia were heavier than Cerrado seeds, but germination of both types was unaffected by submergence. Three-month-old Amazonian and Cerrado seedlings of both species survived 30 days of waterlogging or submersion despite suffering significant inhibition in biomass especially if submerged. Shoot elongation was also arrested. Submersion triggered chlorosis and leaf abscission in Amazonian and Cerrado G. ulmifolia while waterlogging did so only in Cerrado seedlings. During 30 days of re-exposure to non-flooded conditions, G. ulmifolia plants that lost their leaves produced a replacement flush. However, they attained only half the plant dry mass of non-flooded plants. Both submerged and waterlogged G. americana retained their leaves. Consequently, plant dry mass after 30 days recovery was less depressed by these stresses than in G. ulmifolia. Small amounts of cortical aerenchyma were found in roots 2 cm from the tip of well-drained plants. The amount was increased by flooding. Waterlogging but not submergence promoted hypertrophy of lenticels at the stem base of both species and adventitious rooting in G. ulmifolia. Despite some loss of performance in dryland plants, flood tolerance traits were present in wetland and dryland populations of both species. They are part of an overall stress-response potential that permits flexible acclimation to locally flooded conditions.

Expanding our understanding of leaf functional syndromes in savanna systems: the role of plant growth form.

The assessment of leaf strategies has been a common theme in ecology, especially where multiple sources of environmental constraints (fire, seasonal drought, nutrient-poor soils) impose a strong selection pressure towards leaf functional diversity, leading to inevitable tradeoffs among leaf traits, and ultimately to niche segregation among coexisting species. As diversification on leaf functional strategies is dependent on integration at whole plant level, we hypothesized that regardless of phylogenetic relatedness, leaf trait functional syndromes in a multivariate space would be associated with the type of growth form. We measured traits related to leaf gas exchange, structure and nutrient status in 57 coexisting species encompassing all Angiosperms major clades, in a wide array of plant morphologies (trees, shrubs, sub-shrubs, herbs, grasses and palms) in a savanna of Central Brazil. Growth forms differed in mean values for the studied functional leaf traits. We extracted 4 groups of functional typologies: grasses (elevated leaf dark respiration, light-saturated photosynthesis on a leaf mass and area basis, lower values of leaf Ca and Mg), herbs (high values of SLA, leaf N and leaf Fe), palms (high values of stomatal conductance, leaf transpiration and leaf K) and woody eudicots (sub-shrubs, shrubs and trees; low SLA and high leaf Ca and Mg). Despite the large range of variation among species for each individual trait and the independent evolutionary trajectory of individual species, growth forms were strongly associated with particular leaf trait combinations, suggesting clear evolutionary constraints on leaf function for morphologically similar species in savanna ecosystems.

Leaf morphophysiology of a Neotropical mistletoe is shaped by seasonal patterns of host leaf phenology.

Several mistletoe species are able to grow and reproduce on both deciduous and evergreen hosts, suggesting a degree of plasticity in their ability to cope with differences in intrinsic host functions. The aim of this study was to investigate the influence of host phenology on mistletoe water relations and leaf gas exchange. Mistletoe Passovia ovata parasitizing evergreen (Miconia albicans) hosts and P. ovata parasitizing deciduous (Byrsonima verbascifolia) hosts were sampled in a Neotropical savanna. Photosynthetic parameters, diurnal cycles of stomatal conductance, pre-dawn and midday leaf water potential, and stomatal anatomical traits were measured during the peak of the dry and wet seasons, respectively. P. ovata showed distinct water-use strategies that were dependent on host phenology. For P. ovata parasitizing the deciduous host, water use efficiency (WUE; ratio of photosynthetic rate to transpirational water loss) was 2-fold lower in the dry season than in the wet season; in contrast, WUE was maintained at the same level during the wet and dry seasons in P. ovata parasitizing the evergreen host. Generally, mistletoe and host diurnal cycles of stomatal conductance were linked, although there were clear differences in leaf water potential, with mistletoe showing anisohydric behaviour and the host showing isohydric behaviour. Compared to mistletoes attached to evergreen hosts, those parasitizing deciduous hosts had a 1.4-fold lower stomatal density and 1.2-fold wider stomata on both leaf surfaces, suggesting that the latter suffered less intense drought stress. This is the first study to show morphophysiological differences in the same mistletoe species parasitizing hosts of different phenological groups. Our results provide evidence that phenotypical plasticity (anatomical and physiological) might be essential to favour the use of a greater range of hosts.

Anatomia da lâmina foliar de onze espécies lenhosas dominantes nas savanas de Roraima / Leaf anatomy of eleven dominant woody species in the savannas of Roraima

O conhecimento da anatomia da folha é crucial para o entendimento da adaptação das plantas ao ambiente. O objetivo deste estudo foi descrever a anatomia da lâmina foliar de 11 espécies lenhosas, frequentes nas savanas do extremo norte da Amazônia, com ênfase na identificação de atributos adaptativos a ecossistemas abertos, sujeitos a forte insolação e déficit hídrico sazonal. Amostras de folhas foram coletadas e processadas segundo técnicas usuais para estudos de anatomia e histoquímica. Bowdichia virgilioides, Byrsonima coccolobifolia, By. crassifolia, By. verbascifolia, Casearia sylvestris, Curatella americana, Erythroxylum suberosum, Himatanthus articulatus, Miconia albicans, Roupala montana e Xylopia aromatica apresentaram caracteres típicos de plantas heliófilas e xerófilas, como cutícula espessa e estômatos predominantes na face abaxial, além de forte investimento em tecido fotossintético. Em oito das onze espécies, o parênquima paliçádico (PP) ocupa 50% ou mais do espaço do mesofilo. Curatella americana, mesofilo isobilateral, e Bo. virgilioides, mesofilo homogêneo, foram as espécies com maior investimento em PP (~80% e 100%, respectivamente). Além disso, destaca-se a presença de hipoderme (Bo. virgilioides e X. aromatica) ou de epiderme estratificada, densos indumentos, idioblastos cristalíferos e extensões da bainha de feixes. Em síntese, este conjunto de atributos estruturais protege a lâmina foliar contra o excesso de luminosidade, aumenta a resistência mecânica, minimiza a transpiração e contribui para manutenção do balanço hídrico da planta, favorecendo, portanto, o estabelecimento destas espécies nas savanas sazonais do norte da Amazônia.

Knowledge of the anatomical characteristics of the leaf blade is crucial to the understanding of plant adaptation to the environment. The objective of this study was to describe the leaf anatomy of 11 woody species of common occurrence in the open savannas of the northern edge of the Amazon. The focus of the study was on the identification of leaf adaptive features to cope with high irradiances and seasonal water deficits. Leaf samples were fixed and processed by the usual methods for anatomical and histochemical studies. Bowdichia virgilioides, Byrsonima coccolobifolia, By. crassifolia, By. verbascifolia, Casearia sylvestris, Curatella americana, Erythroxylum suberosum, Himatanthus articulatus, Miconia albicans, Roupala montana and Xylopia aromatica showed leaf anatomical traits typical of heliophilous and xerophilous plants such as thick cuticle, stomata prevailing on the abaxial surface, strong investment in photosynthetic tissue. In eight of the eleven species the palisade parenchyma (PP) occupied 50% or more of the mesophyll. Curatella americana, with isobilateral mesophyll, and Bo. virgilioides, with homogeneous mesophyll were the species with the highest investment in PP (~80% and 100%, respectively). Leaves were also characterized by the presence of hypodermis (Bo. virgilioides and X. aromatica) or stratified epidermis, dense indumenta, crystalliferous idioblasts and bundle sheath extensions. This distinctive assortment of anatomical traits helps protecting the leaf blade against excessive irradiances, increases mechanical strength, minimize transpiration and contribute to the maintenance of leaf water balance. Overall they favor the establishment of these species in the seasonal savannas of northern Amazon.

Seed germination and seedling development in response to submergence in tree species of the Central Amazonian floodplains.

Successful germination and seedling establishment are crucial steps for maintenance and expansion of plant populations and recovery from perturbations. Every year the Amazon River and its tributaries overflow and flood the adjacent forest, exerting a strong selective pressure on traits related to seedling recruitment. We examined seed characteristics, stored reserves, germination, seedling development and survival under water of eight representative tree species from the lower portions of the flood-level gradient to identify adaptive strategies that contribute to their regeneration in this extreme ecosystem. Submerged seedlings were assessed for longevity and survival until they showed symptoms of injury. At this point, the remaining healthy seedlings were planted in unsaturated soil to monitor recovery after re-exposure to air over 30 days. All small (seed mass ≤0.17 g) seeds had epigeal phanerocotylar-type germination, a trait that would allow plants to acquire light and CO2 in the shortest time. Cell wall storage polysaccharide was a major component of all seeds, suggesting plant investment in structural reserves. Seven of the eight species germinated and formed healthy seedlings under water that endured submersion without any apparent injury for periods of 20-115 days, depending on the species. Seedlings of some species changed the direction of root growth and grew towards the surface of the water, which might have increased the uptake of oxygen to the tissues. Only one of the seven species did not survive re-exposure to air. Species able to germinate and produce seedlings under submersion, which subsequently are able to establish in aerated soils, would have more time available for terrestrial growth. This is critical for colonization of lower portions of the flood-level gradient where establishment is constrained by the short terrestrial phase that precedes the next flood.

Physiological and proteomic analyses of Saccharum spp. grown under salt stress.

Sugarcane (Saccharum spp.) is the world most productive sugar producing crop, making an understanding of its stress physiology key to increasing both sugar and ethanol production. To understand the behavior and salt tolerance mechanisms of sugarcane, two cultivars commonly used in Brazilian agriculture, RB867515 and RB855536, were submitted to salt stress for 48 days. Physiological parameters including net photosynthesis, water potential, dry root and shoot mass and malondialdehyde (MDA) content of leaves were determined. Control plants of the two cultivars showed similar values for most traits apart from higher root dry mass in RB867515. Both cultivars behaved similarly during salt stress, except for MDA levels for which there was a delay in the response for cultivar RB867515. Analysis of leaf macro- and micronutrients concentrations was performed and the concentration of Mn(2+) increased on day 48 for both cultivars. In parallel, to observe the effects of salt stress on protein levels in leaves of the RB867515 cultivar, two-dimensional gel electrophoresis followed by MS analysis was performed. Four proteins were differentially expressed between control and salt-treated plants. Fructose 1,6-bisphosphate aldolase was down-regulated, a germin-like protein and glyceraldehyde 3-phosphate dehydrogenase showed increased expression levels under salt stress, and heat-shock protein 70 was expressed only in salt-treated plants. These proteins are involved in energy metabolism and defense-related responses and we suggest that they may be involved in protection mechanisms against salt stress in sugarcane.

Plant reproduction in the Central Amazonian floodplains: challenges and adaptations.

BACKGROUND: The Central Amazonian floodplain forests are subjected to extended periods of flooding and to flooding amplitudes of 10 m or more. The predictability, the length of the flood pulse, the abrupt transition in the environmental conditions along topographic gradients on the banks of major rivers in Central Amazonia, and the powerful water and sediment dynamics impose a strong selective pressure on plant reproduction systems. SCOPE: In this review, we examine how the hydrological cycle influences the strategies of sexual and asexual reproduction in herbaceous and woody plants. These are of fundamental importance for the completion of the life cycle. Possible constraints to seed germination, seedling establishment and formation of seed banks are also covered. Likewise, we also discuss the importance of river connectivity for species propagation and persistence in floodplains. CONCLUSIONS: The propagation and establishment strategies employed by the highly diversified assortment of different plant life forms result in contrasting successional stages and a zonation of plant assemblages along the flood-level gradient, whose species composition and successional status are continuously changing not only temporally but also spatially along the river channel.