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Gypsum soils are amongst the most widespread extreme substrates of the world, occurring in 112 countries. This type of hypercalcic substrate has a suite of extreme physical and chemical properties that make it stressful for plant establishment and growth. Extreme chemical properties include low plant-available nitrogen and phosphorus and high plant-available sulphur and calcium, which impose strong nutritional imbalances on plants. In spite of these edaphic barriers, gypsum soils harbour rich endemic floras that have evolved independently on five continents, with highly specialized species. Plants that only grow on gypsum are considered soil specialists, and they have a foliar elemental composition similar to the elemental availability of gypsum soils, with high calcium, sulphur and magnesium accumulation. However, the physiological and ecological role of the unique foliar elemental composition of gypsum specialists remains poorly understood, and it is unknown whether it provides an ecological advantage over other generalist species on gypsum soils. This article reviews available literature on the impact of gypsum soil features on plant life and the mechanisms underlying plant adaptation to gypsum environments. We conclude with a hypothesis on the potential role of the nutritional strategy underlying plant specialization to gypsum soils: Gypsum specialists primarily use SO42- as a counter anion to tolerate high Ca2+ concentrations in cells and avoid phosphorus depletion, which is one of the most limiting nutrients in gypsum soils.
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Extreme soils often have mineral nutrient imbalances compared to plant nutritional requirements and co-occur in open areas where grazers thrive. Thus, plants must respond to both constraints, which can affect nutrient concentrations in all plant organs. Gypsum soil provides an excellent model system to study adaptations to extreme soils under current grazing practices as it harbours two groups of plant species that differ in their tolerance to gypsum soils and foliar composition. However, nutrient concentrations in organs other than leaves, and their individual responses to simulated herbivory, are still unknown in gypsum plants. We studied plant biomass, root mass ratio and nutrient partitioning among different organs (leaves, stems, coarse roots, fine roots) in five gypsum endemics and five generalists cultivated in gypsum and calcareous soils and subjected to different levels of simulated browsing. Gypsum endemics tended to have higher elemental concentration in leaves, stems and coarse roots than generalist species in both soil types, whereas both groups tended to show similar high concentrations in fine roots. This behaviour was especially clear with sulphur (S), which is found in excess in gypsum soils, and which endemics accumulated in leaves as sulphate (>50% of S). Moreover, plants subjected to clipping, regardless of their affinity to gypsum, were unable to compensate for biomass losses and showed similar elemental composition to unclipped plants. The accumulation of excess mineral nutrients by endemic species in aboveground organs may be a constitutive nutritional strategy in extreme soils and is potentially playing an anti-herbivore role in grazed gypsum outcrops.
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Sulfato de Calcio , Suelo , Nutrientes , Hojas de la Planta , Raíces de Plantas , PlantasRESUMEN
The analysis of plant elemental composition and the underlying factors affecting its variation are a current hot topic in ecology. Ecological adaptation to atypical soils may shift plant elemental composition. However, no previous studies have evaluated its relevance against other factors such as phylogeny, climate or individual soil conditions. We evaluated the effect of the phylogeny, environment (climate, soil), and affinity to gypsum soils on the elemental composition of 83 taxa typical of Iberian gypsum ecosystems. We used a new statistical procedure (multiple phylogenetic variance decomposition, MPVD) to decompose total explained variance by different factors across all nodes in the phylogenetic tree of target species (covering 120 million years of Angiosperm evolution). Our results highlight the relevance of phylogeny on the elemental composition of plants both at early (with the development of key preadaptive traits) and recent divergence times (diversification of the Iberian gypsum flora concurrent with Iberian gypsum deposit accumulation). Despite the predominant phylogenetic effect, plant adaptation to gypsum soils had a strong impact on the elemental composition of plants, particularly on sulphur concentrations, while climate and soil effects were smaller. Accordingly, we detected a convergent evolution of gypsum specialists from different lineages on increased sulphur and magnesium foliar concentrations.
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Sulfato de Calcio , Ecosistema , Filogenia , Plantas/genética , Suelo , AzufreRESUMEN
BACKGROUND AND AIMS: Plants depend fundamentally on establishment from seed. However, protocols in trait-based ecology currently estimate seed size but not seed number. This can be rectified. For annuals, seed number should simply be a positive function of vegetative biomass and a negative function of seed size. METHODS: Using published values of comparative seed number as the 'gold standard' and a large functional database, comparative seed yield and number per plant and per m2 were predicted by multiple regression. Subsequently, ecological variation in each was explored for English and Spanish habitats, newly calculated C-S-R strategies and changed abundance in the British flora. KEY RESULTS: As predicted, comparative seed mass yield per plant was consistently a positive function of plant size and competitive ability, and largely independent of seed size. Regressions estimating comparative seed number included, additionally, seed size as a negative function. Relationships differed numerically between regions, habitats and C-S-R strategies. Moreover, some species differed in life history over their geographical range. Comparative seed yield per m2 was positively correlated with FAO crop yield, and increasing British annuals produced numerous seeds. Nevertheless, predicted values must be viewed as comparative rather than absolute: they varied according to the 'gold standard' predictor used. Moreover, regressions estimating comparative seed yield per m2 achieved low precision. CONCLUSIONS: For the first time, estimates of comparative seed yield and number for >800 annuals and their predictor equations have been produced and the ecological importance of these regenerative traits has been illustrated. 'Regenerative trait-based ecology' remains in its infancy, with work needed on determinate vs. indeterminate flowering ('bet-hedging'), C-S-R methodologies, phylogeny, comparative seed yield per m2 and changing life history. Nevertheless, this has been a positive start and readers are invited to use estimates for >800 annuals, in the Supplementary data, to help advance 'regenerative trait-based ecology' to the next level.
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Plantas , Semillas , Ecosistema , Fenotipo , FilogeniaRESUMEN
Background and Aims: While the 'worldwide leaf economics spectrum' (Wright IJ, Reich PB, Westoby M, et al. 2004. The worldwide leaf economics spectrum. Nature : 821-827) defines mineral nutrient relationships in plants, no unifying functional consensus links size attributes. Here, the focus is upon leaf size, a much-studied plant trait that scales positively with habitat quality and components of plant size. The objective is to show that this wide range of relationships is explicable in terms of a seed-phytomer-leaf (SPL) theoretical model defining leaf size in terms of trade-offs involving the size, growth rate and number of the building blocks (phytomers) of which the young shoot is constructed. Methods: Functional data for 2400+ species and English and Spanish vegetation surveys were used to explore interrelationships between leaf area, leaf width, canopy height, seed mass and leaf dry matter content (LDMC). Key Results: Leaf area was a consistent function of canopy height, LDMC and seed mass. Additionally, size traits are partially uncoupled. First, broad laminas help confer competitive exclusion while morphologically large leaves can, through dissection, be functionally small. Secondly, leaf size scales positively with plant size but many of the largest-leaved species are of medium height with basally supported leaves. Thirdly, photosynthetic stems may represent a functionally viable alternative to 'small seeds + large leaves' in disturbed, fertile habitats and 'large seeds + small leaves' in infertile ones. Conclusions: Although key elements defining the juvenile growth phase remain unmeasured, our results broadly support SPL theory in that phytometer and leaf size are a product of the size of the initial shoot meristem (â seed mass) and the duration and quality of juvenile growth. These allometrically constrained traits combine to confer ecological specialization on individual species. Equally, they appear conservatively expressed within major taxa. Thus, 'evolutionary canalization' sensu Stebbins (Stebbins GL. 1974. Flowering plants: evolution above the species level . Cambridge, MA: Belknap Press) is perhaps associated with both seed and leaf development, and major taxa appear routinely specialized with respect to ecologically important size-related traits.
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Rasgos de la Historia de Vida , Hojas de la Planta/fisiología , Fenómenos Fisiológicos de las Plantas , Semillas/fisiología , Ecosistema , Inglaterra , Hojas de la Planta/anatomía & histología , Hojas de la Planta/crecimiento & desarrollo , Semillas/anatomía & histología , Semillas/crecimiento & desarrollo , SueciaRESUMEN
Gypsum soils are among the most restrictive and widespread substrates for plant life. Plants living on gypsum are classified as gypsophiles (exclusive to gypsum) and gypsovags (non-exclusive to gypsum). The former have been separated into wide and narrow gypsophiles, each with a putative different ecological strategy. Mechanisms displayed by gypsum plants to compete and survive on gypsum are still not fully understood. The aim of this study was to compare the main chemical groups in the leaves of plants with different specificity to gypsum soils and to explore the ability of Fourier transform infrared (FTIR) spectra analyzed with neural network (NN) modelling to discriminate groups of gypsum plants. Leaf samples of 14 species with different specificity to gypsum soils were analysed with FTIR spectroscopy coupled to neural network (NN) modelling. Spectral data were further related to the N, C, S, P, K, Na, Ca, Mg and ash concentrations of samples. The FTIR spectra of the three groups analyzed showed distinct features that enabled their discrimination through NN models. Wide gypsophiles stood out for the strong presence of inorganic compounds in their leaves, particularly gypsum and, in some species, also calcium oxalate crystals. The spectra of gypsovags had less inorganic chemical species, while those of narrow gypsum endemisms had low inorganics but shared with wide gypsophiles the presence of oxalate. Gypsum and calcium oxalate crystals seem to be widespread amongst gypsum specialist plants, possibly as a way to tolerate excess Ca and sulphate. However, other mechanisms such as the accumulation of sulphates in organic molecules are also compatible with plant specialization to gypsum. While gypsovags seem to be stress tolerant plants that tightly regulate the uptake of S and Ca, the ability of narrow gypsum endemisms to accumulate excess Ca as oxalate may indicate their incipient specialization to gypsum.
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Plantas/química , Suelo/química , Espectroscopía Infrarroja por Transformada de Fourier/métodos , Elementos Químicos , Hojas de la Planta/químicaRESUMEN
Some minerals, like gypsum, hold water in their crystalline structure. Although still unexplored, the use of such crystallization water by organisms would point to a completely new water source for life, critical under dry conditions. Here we use the fact that the isotopic composition of free water differs from gypsum crystallization water to show that plants can use crystallization water from the gypsum structure. The composition of the xylem sap of gypsum plants during summer shows closer values to gypsum crystallization water than to free soil water. Crystallization water represents a significant water source for organisms growing on gypsum, especially during summer, when it accounts for 70-90% of the water used by shallow-rooted plants. Given the widespread occurrence of gypsum in dry lands throughout the Earth and in Mars, these results may have important implications for arid land reclamation and exobiology.
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Sulfato de Calcio/química , Fenómenos Fisiológicos de las Plantas , Agua/análisis , Teorema de Bayes , Cristalización , Exobiología , Xilema/fisiologíaRESUMEN
Nitrogen (N) is, after water, the most limiting resource in semiarid ecosystems. However, knowledge on the N cycling ability of semiarid woody plants is still very rudimentary. This study analyzed the seasonal change in the N concentrations and pools of the leaves and woody organs of two species of semiarid sub-shrubs with contrasting leaf habit. The ability of both species to uptake, remobilize and recycle N, plus the main storage organ for N during summer drought were evaluated. We combined an observational approach in the field with experimental (15)N labelling of adult individuals grown in sand culture. Seasonal patterns of N concentrations were different between species and organs and foliar N concentrations of the summer deciduous Lepidium subulatum were almost double those of the evergreen Linum suffruticosum. L. subulatum up took ca. 60% more external N than the evergreen and it also had a higher N resorption efficiency and proficiency. Contrastingly, L. suffruticosum relied more on internal N remobilization for shoot growth. Differently to temperate species, the evergreen stored N preferentially in the main stem and old trunks, while the summer deciduous stored it in the foliage and young stems. The higher ability of L. subulatum to uptake external N can be related to its ability to perform opportunistic growth and exploit the sporadic pulses of N typical of semiarid ecosystems. Such ability may also explain its high foliar N concentrations and its preferential storage of N in leaves and young stems. Finally, L. suffruticosum had a lower ability to recycle N during leaf senescence. These strategies contrast with those of evergreen and deciduous species from temperate and boreal areas, highlighting the need of further studies on semiarid and arid plants.
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Lepidium/metabolismo , Ciclo del Nitrógeno , Hojas de la Planta/metabolismo , Análisis de Varianza , Fenotipo , Brotes de la Planta/metabolismo , Estaciones del AñoRESUMEN
BACKGROUND AND AIMS: In trees, bud development is driven by endogenous and exogenous factors such as species and climate, respectively. However, knowledge is scarce on how these factors drive changes in bud size across different time scales. METHODS: The seasonal patterns of apical bud enlargement are related to primary and secondary growth in two coexisting Mediterranean oaks with contrasting leaf habit (Quercus ilex, evergreen; Quercus faginea, deciduous) over three years. In addition, the climatic factors driving changes in bud size of the two oak species were determined by correlating bud mass with climatic variables at different time scales (from 5 to 30 d) over a 15-year period. KEY RESULTS: The maximum enlargement rate of buds was reached between late July and mid-August in both species. Moreover, apical bud size increased with minimum air temperatures during the period of maximum bud enlargement rates. CONCLUSIONS: The forecasted rising minimum air temperatures predicted by climatic models may affect bud size and consequently alter crown architecture differentially in sympatric Mediterranean oaks. However, the involvement of several drivers controlling the final size of buds makes it difficult to predict the changes in bud size as related to ongoing climate warming.
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Clima , Quercus/crecimiento & desarrollo , Hojas de la Planta/crecimiento & desarrollo , Brotes de la Planta/crecimiento & desarrollo , Tallos de la Planta/crecimiento & desarrollo , Estaciones del Año , España , Temperatura , ÁrbolesRESUMEN
BACKGROUND AND AIMS: Plant species typical of cold and warm habitats differ in a suite of morpho-physio-phenological traits, although their evolutionary routes have been poorly explored. Here, it is advocated that traits typical of different climate regimes can be largely driven by contrasting branch architectures. This is explored within Saxifraga. First, an investigation was carried out to determine whether series Ceratophyllae (lateral inflorescences) is segregated to lowlands compared with Pentadactylis (terminal inflorescences). Then, two altitudinal vicariants, S. trifurcata (lowland, with lateral inflorescences) and S. canaliculata (highland, with apical inflorescences), were selected. It was hypothesized that apical flowering of S. canaliculata constrains its growth period, bringing with it traits typical of short growth season plants, and conversely for S. trifurcata. METHODS: The hypothesis was tested by measuring plant compactness and organ pre-formation in seven populations of these species along an altitude gradient. KEY RESULTS: Most variables differed among species. Morphological variables at all scales support that the architecture of S. canaliculata generates a more compact habit. A higher number of primordia and earlier inflorescence pre-formation in S. canaliculata indicate that it begins organogenesis earlier. Data on organogenesis suggest that the different timing of inflorescence initiation may be the origin of the contrasting architectures. Within species, shoot compactness increased, and the length of lateral primordia decreased, as altitude increased. All other metrics were similar among locations of the same species at contrasting altitudes. CONCLUSIONS: The hypotheses linking elevational segregation of species, architecture and pheno-morphological traits were validated at broad (gen. Saxifraga) and local (altitudinal vicariants) scales. This supports the initial idea that shoot architecture may to a large extent condition high altitude adaptive syndrome.
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Altitud , Brotes de la Planta/anatomía & histología , Saxifragaceae/anatomía & histología , Modelos Biológicos , Hojas de la Planta/anatomía & histología , Carácter Cuantitativo Heredable , Estaciones del Año , Especificidad de la Especie , Factores de TiempoRESUMEN
This study assesses how different phases of shoot growth underlie seasonal change in leaf and stem dry matter content (LDMC and SDMC, respectively) of 12 woody Mediterranean species. The relationship between LDMC and nonstructural carbohydrate (NSC) concentrations is also explored and the seasonal vs interspecies variability of LDMC compared. LDMC, SDMC and shoot elongation rate (SER) were measured on a monthly basis for a minimum of 12 months. Bud growth rate (BGR) and NSC concentrations were also assessed in several of the study species. LDMC and SDMC decreased during shoot elongation in spring and increased in summer, showing a significant negative correlation with SER, but were unrelated to BGR. Half of the species analysed showed a positive relationship between LDMC and NSC. Seasonal fluctuations of LDMC within species were higher than interspecies differences, and species ranking was significantly affected by the month of sampling, except during winter months. Seasonal changes in LDMC and SDMC are mainly related to shoot elongation phenology, and NSC sink-source relationships between old and growing organs can explain this relationship in some species. Owing to the high seasonal variability in LDMC, it is recommended that samples for comparative purposes should be collected as close to the winter as possible.
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Metabolismo de los Hidratos de Carbono , Plantas/metabolismo , Estaciones del Año , Desarrollo de la Planta , Hojas de la Planta/crecimiento & desarrollo , Hojas de la Planta/metabolismo , Brotes de la Planta/crecimiento & desarrollo , Brotes de la Planta/metabolismo , Tallos de la Planta/crecimiento & desarrollo , Tallos de la Planta/metabolismo , Especificidad de la EspecieRESUMEN
BACKGROUND AND AIMS: This study analysed the differences in nitrogen (N), non-structural carbohydrates (NSC) and biomass allocation to the roots and shoots of 18 species of Mediterranean dwarf shrubs with different shoot-rooting and resprouting abilities. Root N and NSC concentrations of strict root-sprouters and species resprouting from the base of the stems were also compared. METHODS: Soluble sugars (SS), starch and N concentrations were assessed in roots and shoots. The root : shoot ratio of each species was obtained by thorough root excavations. Cross-species analyses were complemented by phylogenetically independent contrasts (PICs). KEY RESULTS: Shoot-rooting species showed a preferential allocation of starch to shoots rather than roots as compared with non-shoot-rooting species. Resprouters displayed greater starch concentrations than non-sprouters in both shoots and roots. Trends were maintained after PICs analyses, but differences became weak when root-sprouters versus non-root-sprouters were compared. Within resprouters, strict root-sprouters showed greater root concentrations and a preferential allocation of starch to the roots than stem-sprouters. No differences were found in the root : shoot ratio of species with different rooting and resprouting abilities. CONCLUSIONS: The shoot-rooting ability of Mediterranean dwarf shrubs seems to depend on the preferential allocation of starch and SS to shoots, though alternative C-sources such as current photosynthates may also be involved. In contrast to plants from other mediterranean areas of the world, the resprouting ability of Mediterranean dwarf shrubs is not related to a preferential allocation of N, NSC and biomass to roots.
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Metabolismo de los Hidratos de Carbono , Nitrógeno/metabolismo , Plantas/metabolismo , Biomasa , Desarrollo de la Planta , Raíces de Plantas/crecimiento & desarrollo , Raíces de Plantas/metabolismo , Brotes de la Planta/crecimiento & desarrollo , Brotes de la Planta/metabolismo , España , Especificidad de la EspecieRESUMEN
BACKGROUND AND AIMS: Plants from gypsum habitats are classified as gypsophiles and gypsovags. The former include both narrow endemics limited to small gypsum areas and regionally dominant gypsophiles growing in gypsum areas of large regions, whereas gypsovags are plants that can grow both in gypsum and non-gypsum soils. Factors controlling the distribution of gypsum plants are still not fully understood. METHODS: To assess how the different types of gypsum plants deal with the stressful conditions of gypsum substrates, comparisons were made of the leaf chemical composition of four gypsovags, five regionally dominant gypsophiles and four narrow gypsum endemics growing in two massive gypsum areas of the Iberian Peninsula. KEY RESULTS: The chemical composition of gypsovags was clearly different from regionally dominant gypsophiles, while the chemical composition of narrow-gypsophile endemics was more similar to the chemical composition of gypsovags than to that of regionally dominant gypsophiles. Regionally dominant gypsophiles showed higher concentrations of ash, Ca, S, N, Mg P and Na, whereas gypsovags and local gypsophile endemics displayed higher concentrations of C and greater C : N ratios. CONCLUSIONS: Such differences suggest that the three groups of gypsum plants follow diverse ecological strategies. It is suggested that regionally dominant gypsophiles might fit the 'specialist' model, being species specifically adapted to gypsum, whereas both gypsovags and narrow-gypsophile endemics might fit the 'refuge' model, being stress-tolerant species that find refuge on gypsum soils from competition. The analysis of the leaf chemical composition could be a good predictor of the degree of plants specialization to gypsum soils.
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Sulfato de Calcio/química , Sulfato de Calcio/metabolismo , Plantas/química , Plantas/metabolismo , Suelo/análisis , Calcio/análisis , Calcio/metabolismo , Magnesio/análisis , Magnesio/metabolismo , EspañaRESUMEN
BACKGROUND AND AIMS: The objective of this study was to test whether the mean values of several root morphological variables were related to the ability to develop root-borne shoots and/or shoot-borne roots in a wide range of vascular plants. METHODS: A comparative study was carried out on the 123 most common plant species from eroded lands in north-east Spain. After careful excavations in the field, measurements were taken of the maximum root depth, absolute and relative basal root diameter, specific root length (SRL), and the root depth/root lateral spread ratio on at least three individuals per species. Shoot-rooting and root-sprouting were observed in a large number of individuals in many eroded and sedimentary environments. The effect of life history and phylogeny on shoot-rooting and root-sprouting abilities was also analysed. KEY RESULTS: The species with coarse and deep tap-roots tended to be root-sprouting and those with fine, fasciculate and long main roots (which generally spread laterally), tended to be shoot-rooting. Phylogeny had an important influence on root system morphology and shoot-rooting and root-sprouting capacities. However, the above relations stood after applying analyses based on phylogenetically independent contrasts (PICs). CONCLUSIONS: The main morphological features of the root system of the study species are related to their ability to sprout from their roots and form roots from their shoots. According to the results, such abilities might only be functionally viable in restricted root system morphologies and ecological strategies.
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Ambiente , Raíces de Plantas/crecimiento & desarrollo , Brotes de la Planta/crecimiento & desarrollo , Filogenia , Desarrollo de la Planta , Raíces de Plantas/anatomía & histología , Raíces de Plantas/clasificación , Brotes de la Planta/anatomía & histología , Brotes de la Planta/clasificación , Plantas/anatomía & histología , Plantas/clasificación , EspañaRESUMEN
BACKGROUND AND AIMS: Understanding the effects of the environment on the morphology and shoot growth activities of plants is crucial to identifying plant ecological strategies. This study analysed the bud morphology, bud activity, shoot growth dynamics and shoot water content at full hydration (WC(h)) of two species of Mediterranean sub-shrubs, Lepidium subulatum and Linum suffruticosum, co-existing in gypsum outcrops in north-east Spain. METHODS: Sampling was conducted monthly over 2 years in one population per species. Buds were dissected under a stereo-microscope. Shoot growth was measured as the mean increase in shoot length of 15 marked individuals between two consecutive samplings. Bud activity was studied following the variations in the number of leaf primordia shorter than 1 mm and longer than 0.025 mm in the buds. KEY RESULTS: Both species bore naked buds and displayed discontinuous seasonal patterns of shoot growth, leaf primordia formation and WC(h). The number of leaf primordia in the bud peaked before the beginning of shoot expansion. In both species, organogenesis and expansion were uncoupled throughout the year. The time lapse between these two processes varied throughout the year, and was greatest for those elements differentiated in autumn. WC(h) was more closely related to shoot expansion than to organogenesis. CONCLUSIONS: Both species displayed similar bud morphology and similar seasonal patterns of bud and shoot growth, and WC(h) as a result of the strong seasonality of the Mediterranean climate in gypsum outcrops. The beginning of the spring period of expansion of long branches coincided with maximum values of WC(h), while the rest period of summer matched minimum values. These results support the hypothesis that the growth of long branches is strongly related to WC(h).