Plant Foliar Geometry as a Biomimetic Template for Antenna Design.

Plant diversity includes over 300,000 species, and leaf structure is one of the main targets of selection, being highly variable in shape and size. On the other hand, the optimization of antenna design has no unique solution to satisfy the current range of applications. We analyzed the foliar geometries of 100 plant species and applied them as a biomimetic design template for microstrip patch antenna systems. From this set, a subset of seven species were further analyzed, including species from tropical and temperate forests across the phylogeny of the Angiosperms. Foliar geometry per species was processed by image processing analyses, and the resultant geometries were used in simulations of the reflection coefficients and the radiation patterns via finite differences methods. A value below -10 dB is set for the reflection coefficient to determine the operation frequencies of all antenna elements. All species showed between 3 and 15 operational frequencies, and four species had operational frequencies that included the 2.4 and 5 GHz bands. The reflection coefficients and the radiation patterns in most of the designs were equal or superior to those of conventional antennas, with several species showing multiband effects and omnidirectional radiation. We demonstrate that plant structures can be used as a biomimetic tool in designing microstrip antenna for a wide range of applications.

Leaf habit determines the hydraulic and resource-use strategies in tree saplings from the Sonoran Desert.

The drought susceptibility of woody saplings may explain their low survival in arid environments. Therefore, it is critical to determine which morphological and physiological traits are more responsive to drought among young plants. This study tested whether plant responses to experimental drought differ between two plant functional groups: the deciduous and evergreen species. We predicted that deciduous species would present a tighter stomatal control under drought, coupled with fast carbon fixation under no stress, tending toward isohydry and faster growth rates than the evergreen species. Using 1-year-old saplings from three evergreen and four deciduous Sonoran Desert tree species, we evaluated their hydraulic and gas exchange traits under three experimental irrigation conditions: high, intermediate and low water availability. We measured CO2 assimilation rates (A), stomatal conductance (gs), the level of iso-anisohydry (as the plant's ability to maintain constant their water potential) and seven morphological and growth-related traits throughout 2 months. Under high water availability, saplings reached their maximum values of A and gs, which were significantly higher for deciduous than evergreen species. Correlations among hydroscape area (HA) and leaf traits positioned species along the iso/anisohydric continuum. Deciduous species presented isohydric characteristics, including low HA, high gs, A and Huber values (HVs), and traits indicative of a faster use of resources, such as low stem-specific density (SSD) and low leaf mass per area (LMA). By contrast, evergreen species showed traits that indicate slow resource use and anisohydric behavior, such as high HA, SSD and LMA, and low gs, A and HVs. Deciduous species drastically reduced gas exchange rates in response to drought, while evergreen maintained low rates independently of drought intensity. Overall, desert saplings showed strategies concordant with the iso-anisohydric continuum and the fast-slow use of resources.

Microstructure and Hydrophobicity of the External Surface of a Sonoran Desert Beetle.

We have studied the external surface (elytra) of the Sonoran Desert beetle (Eleodes eschscholtzii). Our aim was to assess whether this species has similar traits to some beetles from the Namibian Desert that are known to have hierarchical micropatterns that allow for water harvesting. We have conducted scanning electron microscopy (SEM) and apparent contact angle experiments on specimens collected at two sampling sites with different ambient humidity. The results show that the beetle's external surface microstructure is composed of a compact array of polygons with randomly scattered protuberances. The density of the polygons in the microstructure is different for individuals collected in different sites: the polygon array is denser in the more humid site and less dense in the drier site. The measured contact angles also depend on the sampling site. For individuals collected in the drier site, the average apparent contact angle is 70°, whereas for the more humid site, the average apparent contact angle is 92°. FT-IR experiments are consistent with the presence of hydrophobic wax compounds in the studied surfaces. Our investigation opens new questions that are currently being addressed by experiments that are underway. For instance, it would be interesting to know whether the observed nanopatterns could be used in biomimetic devices for water harvesting purposes.

Hydrological and topographic determinants of biomass and species richness in a Mediterranean-climate shrubland.

BACKGROUND: In arid and semiarid shrublands, water availability directly influences ecosystem properties. However, few empirical tests have determined the association between particular soil and hydrology traits with biodiversity and ecosystem biomass at the local scale. METHODS: We tested if plant species richness (S) and aboveground biomass (AGB) were associated with soil and topographic properties on 36 plots (ca. 12.5 m2) in 17 hectares of chaparral in the Mediterranean-climate of Valle de Guadalupe, Baja California, México. We used close-to-the-ground aerial photography to quantify sky-view cover per species, including all growth forms. We derived an elevation model (5 cm) from other aerial imagery. We estimated six soil properties (soil water potential, organic matter content, water content, pH, total dissolved solids concentration, and texture) and four landscape metrics (slope, aspect, elevation, and topographic index) for the 36 plots. We quantified the biomass of stems, leaves, and reproductive structures, per species. RESULTS: 86% of AGB was in stems, while non-woody species represented 0.7% of AGB but comprised 38% of S (29 species). Aboveground biomass and species richness were unrelated across the landscape. S was correlated with aspect and elevation (R = 0.53, aspect P = 0.035, elevation P = 0.05), while AGB (0.006-9.17 Kg m-2) increased with soil water potential and clay content (R = 0.51, P = 0.02, and P = 0.04). Only three species (11% of total S) occupied 65% of the total plant cover, and the remaining 26 represented only 35%. Cover was negatively correlated with S (R = -0.38, P = 0.02). 75% of AGB was concentrated in 30% of the 36 plots, and 96% of AGB corresponded to only 20% of 29 species. DISCUSSION: At the scale of small plots in our studied Mediterranean-climate shrubland in Baja California, AGB was most affected by soil water storage. AGB and cover were dominated by a few species, and only cover was negatively related to S. S was comprised mostly by uncommon species and tended to increase as plant cover decreased.

Leaf water relations reflect canopy phenology rather than leaf life span in Sonoran Desert trees.

Plants from arid environments display covarying traits to survive or resist drought. Plant drought resistance and ability to survive long periods of low soil water availability should involve leaf phenology coordination with leaf and stem functional traits related to water status. This study tested correlations between phenology and functional traits involved in plant water status regulation in 10 Sonoran Desert tree species with contrasting phenology. Species seasonal variation in plant water status was defined by calculating their relative positions along the iso/anisohydric regulation continuum based on their hydroscape areas (HA)-a metric derived from the relationship between predawn and midday water potentials-and stomatal and hydraulic traits. Additionally, functional traits associated with plant water status regulation, including lamina vessel hydraulic diameter (DHL), stem-specific density (SSD) and leaf mass per area (LMA) were quantified per species. To characterize leaf phenology, leaf longevity (LL) and canopy foliage duration (FD) were determined. Hydroscape area was strongly correlated with FD but not with leaf longevity (LL); HA was significantly associated with SSD and leaf hydraulic traits (DHL, LMA) but not with stem hydraulic traits (vulnerability index, relative conductivity); and FD was strongly correlated with LMA and SSD. Leaf physiological characteristics affected leaf phenology when it was described as canopy FD better than when described as LL. Stem and leaf structure and hydraulic functions were not only relevant for categorizing species along the iso/anisohydric continuum but also allowed identifying different strategies of desert trees within the 'fast-slow' plant economics spectrum. The results in this study pinpoint the set of evolutionary pressures that shape the Sonoran Desert Scrub physiognomy.

Plant height and hydraulic vulnerability to drought and cold.

Understanding how plants survive drought and cold is increasingly important as plants worldwide experience dieback with drought in moist places and grow taller with warming in cold ones. Crucial in plant climate adaptation are the diameters of water-transporting conduits. Sampling 537 species across climate zones dominated by angiosperms, we find that plant size is unambiguously the main driver of conduit diameter variation. And because taller plants have wider conduits, and wider conduits within species are more vulnerable to conduction-blocking embolisms, taller conspecifics should be more vulnerable than shorter ones, a prediction we confirm with a plantation experiment. As a result, maximum plant size should be short under drought and cold, which cause embolism, or increase if these pressures relax. That conduit diameter and embolism vulnerability are inseparably related to plant size helps explain why factors that interact with conduit diameter, such as drought or warming, are altering plant heights worldwide.

Osmotic and hydraulic adjustment of mangrove saplings to extreme salinity.

Salinity tolerance in plant species varies widely due to adaptation and acclimation processes at the cellular and whole-plant scales. In mangroves, extreme substrate salinity induces hydraulic failure and ion excess toxicity and reduces growth and survival, thus suggesting a potentially critical role for physiological acclimation to salinity. We tested the hypothesis that osmotic adjustment, a key type of plasticity that mitigates salinity shock, would take place in coordination with declines in whole-plant hydraulic conductance in a common garden experiment using saplings of three mangrove species with different salinity tolerances (Avicennia germinans L., Rhizophora mangle L. and Laguncularia racemosa (L.) C.F. Gaertn., ordered from higher to lower salinity tolerance). For each mangrove species, four salinity treatments (1, 10, 30 and 50 practical salinity units) were established and the time trajectories were determined for leaf osmotic potential (Ψs), stomatal conductance (gs), whole-plant hydraulic conductance (Kplant) and predawn disequilibrium between xylem and substrate water potentials (Ψpdd). We expected that, for all three species, salinity increments would result in coordinated declines in Ψs, gs and Kplant, and that the Ψpdd would increase with substrate salinity and time of exposure. In concordance with our predictions, reductions in substrate water potential promoted a coordinated decline in Ψs, gs and Kplant, whereas the Ψpdd increased substantially during the first 4 days but dissipated after 7 days, indicating a time lag for equilibration after a change in substratum salinity. Our results show that mangroves confront and partially ameliorate acute salinity stress via simultaneous reductions in Ψs, gs and Kplant, thus developing synergistic physiological responses at the cell and whole-plant scales.

Root biomechanics in Rhizophora mangle: anatomy, morphology and ecology of mangrove's flying buttresses.

BACKGROUND AND AIMS: Rhizophora species of mangroves have a conspicuous system of stilt-like roots (rhizophores) that grow from the main stem and resemble flying buttresses. As such, the development of rhizophores can be predicted to be important for the effective transmission of dynamic loads from the top of the tree to the ground, especially where the substrate is unstable, as is often the case in the habitats where Rhizophora species typically grow. This study tests the hypothesis that rhizophore architecture in R. mangle co-varies with their proximity to the main stem, and with stem size and crown position. METHODS: The allometry and wood mechanical properties of R. mangle (red mangrove) trees growing in a mangrove basin forest within a coastal lagoon in Mexico were compared with those of coexisting, non-buttressed mangrove trees of Avicennia germinans. The anatomy of rhizophores was related to mechanical stress due to crown orientation (static load) and to prevailing winds (dynamic load) at the study site. KEY RESULTS: Rhizophores buttressed between 10 and 33 % of tree height. There were significant and direct scaling relationships between the number, height and length of rhizophores vs. basal area, tree height and crown area. Wood mechanical resistance was significantly higher in the buttressed R. mangle (modulus of elasticity, MOE = 18·1 ± 2 GPa) than in A. germinans (MOE = 12·1 ± 0·5 GPa). Slenderness ratios (total height/stem diameter) were higher in R. mangle, but there were no interspecies differences in critical buckling height. When in proximity to the main stem, rhizophores had a lower length/height ratio, higher eccentricity and higher xylem/bark and pith proportions. However, there were no directional trends with regard to prevailing winds or tree leaning. CONCLUSIONS: In comparison with A. germinans, a tree species with wide girth and flare at the base, R. mangle supports a thinner stem of higher mechanical resistance that is stabilized by rhizophores resembling flying buttresses. This provides a unique strategy to increase tree slenderness and height in the typically unstable substrate on which the trees grow, at a site that is subject to frequent storms.

Leaf mass per area is independent of vein length per area: avoiding pitfalls when modelling phenotypic integration (reply to Blonder et al. 2014).

It has been recently proposed that leaf vein length per area (VLA) is the major determinant of leaf mass per area ( MA), and would thereby determine other traits of the leaf economic spectrum (LES), such as photosynthetic rate per mass (A(mass)), nitrogen concentration per mass (N(mass)) and leaf lifespan (LL). In a previous paper we argued that this 'vein origin' hypothesis was supported only by a mathematical model with predestined outcomes, and that we found no support for the 'vein origin' hypothesis in our analyses of compiled data. In contrast to the 'vein origin' hypothesis, empirical evidence indicated that VLA and LMA are independent mechanistically, and VLA (among other vein traits) contributes to a higher photosynthetic rate per area (A(area)), which scales up to driving a higher A(mass), all independently of LMA, N(mass) and LL. In their reply to our paper, Blonder et al. (2014) raised questions about our analysis of their model, but did not address our main point, that the data did not support their hypothesis. In this paper we provide further analysis of an extended data set, which again robustly demonstrates the mechanistic independence of LMA from VLA, and thus does not support the 'vein origin' hypothesis. We also address the four specific points raised by Blonder et al. (2014) regarding our analyses. We additionally show how this debate provides critical guidance for improved modelling of LES traits and other networks of phenotypic traits that determine plant performance under contrasting environments.

Dynamic control of osmolality and ionic composition of the xylem sap in two mangrove species.

• Premise of the study: Xylem sap osmolality and salinity is a critical unresolved issue in plant function with impacts on transport efficiency, pressure gradients, and living cell turgor pressure, especially for halophytes such as mangrove trees.• Methods: We collected successive xylem vessel sap samples from stems and shoots of Avicennia germinans and Laguncularia racemosa using vacuum and pressure extraction and measured their osmolality. Following a series of extractions with the pressure chamber, we depressurized the shoot and pressurized again after various equilibration periods (minutes to hours) to test for dynamic control of osmolality. Transpiration and final sap osmolality were measured in shoots perfused with deionized water or different seawater dilutions.• Key results: For both species, the sap osmolality values of consecutive samples collected by vacuum extraction were stable and matched those of the initial samples extracted with the pressure chamber. Further extraction of samples with the pressure chamber decreased sap osmolality, suggesting reverse osmosis occurred. However, sap osmolalities increased when longer equilibration periods after sap extraction were allowed. Analysis of expressed sap with HPLC indicated a 1:1 relation between measured osmolality and the osmolality of the inorganic ions in the sap (mainly Na+, K+, and Cl-), suggesting no contamination by organic compounds. In stems perfused with deionized water, the sap osmolality increased to mimic the native sap osmolality.• Conclusions: Xylem sap osmolality and ionic contents are dynamically adjusted by mangroves and may help modulate turgor pressure, hydraulic conductivity, and water potential, thus being important for mangrove physiology, survival, and distribution.

Specific polyphenols and tannins are associated with defense against insect herbivores in the tropical oak Quercus oleoides.

The role of plant polyphenols as defenses against insect herbivores is controversial. We combined correlative field studies across three geographic regions (Northern Mexico, Southern Mexico, and Costa Rica) with induction experiments under controlled conditions to search for candidate compounds that might play a defensive role in the foliage of the tropical oak, Quercus oleoides. We quantified leaf damage caused by four herbivore guilds (chewers, skeletonizers, leaf miners, and gall forming insects) and analyzed the content of 18 polyphenols (including hydrolyzable tannins, flavan-3-ols, and flavonol glycosides) in the same set of leaves using high performance liquid chromatography and mass spectrometry. Foliar damage ranged from two to eight percent per region, and nearly 90% of all the damage was caused by chewing herbivores. Damage due to chewing herbivores was positively correlated with acutissimin B, catechin, and catechin dimer, and damage by mining herbivores was positively correlated with mongolinin A. By contrast, gall presence was negatively correlated with vescalagin and acutissimin B. By using redundancy analysis, we searched for the combinations of polyphenols that were associated to natural herbivory: the combination of mongolinin A and acutissimin B had the highest association to herbivory. In a common garden experiment with oak saplings, artificial damage increased the content of acutissimin B, mongolinin A, and vescalagin, whereas the content of catechin decreased. Specific polyphenols, either individually or in combination, rather than total polyphenols, were associated with standing leaf damage in this tropical oak. Future studies aimed at understanding the ecological role of polyphenols can use similar correlative studies to identify candidate compounds that could be used individually and in biologically meaningful combinations in tests with herbivores and pathogens.

Bark functional ecology: evidence for tradeoffs, functional coordination, and environment producing bark diversity.

The causes underlying bark diversity are unclear. Variation has been frequently attributed to environmental differences across sites. However, variation may also result from tradeoffs and coordination between bark's multiple functions. Bark traits may also covary with wood and leaf traits as part of major dimensions of plant variation. To assess hypotheses regarding tradeoffs and functional coordination, we measured bark traits reflecting protection, storage, mechanics, and photosynthesis in branches of 90 species spanning a wide phylogenetic and environmental range. We also tested associations between bark, wood, and leaf traits. We partitioned trait variation within species, and within and across communities to quantify variation associated with across-site differences. We observed associations between bark mechanics and storage, density and thickness, and thickness and photosynthetic activity. Increasing bark thickness contributed significantly to stiffer stems and greater water storage. Bark density, water content, and mechanics covaried strongly with the equivalent wood traits, and to a lesser degree with leaf size, xylem conductivity, and vessel diameter. Most variation was observed within sites and had low phylogenetic signal. Compared with relatively minor across-site differences, tradeoffs and coordination among functions of bark, leaves, and wood are likely to be major and overlooked factors shaping bark ecology and evolution.

How do leaf veins influence the worldwide leaf economic spectrum? Review and synthesis.

Leaf vein traits are implicated in the determination of gas exchange rates and plant performance. These traits are increasingly considered as causal factors affecting the 'leaf economic spectrum' (LES), which includes the light-saturated rate of photosynthesis, dark respiration, foliar nitrogen concentration, leaf dry mass per area (LMA) and leaf longevity. This article reviews the support for two contrasting hypotheses regarding a key vein trait, vein length per unit leaf area (VLA). Recently, Blonder et al. (2011, 2013) proposed that vein traits, including VLA, can be described as the 'origin' of the LES by structurally determining LMA and leaf thickness, and thereby vein traits would predict LES traits according to specific equations. Careful re-examination of leaf anatomy, published datasets, and a newly compiled global database for diverse species did not support the 'vein origin' hypothesis, and moreover showed that the apparent power of those equations to predict LES traits arose from circularity. This review provides a 'flux trait network' hypothesis for the effects of vein traits on the LES and on plant performance, based on a synthesis of the previous literature. According to this hypothesis, VLA, while virtually independent of LMA, strongly influences hydraulic conductance, and thus stomatal conductance and photosynthetic rate. We also review (i) the specific physiological roles of VLA; (ii) the role of leaf major veins in influencing LES traits; and (iii) the role of VLA in determining photosynthetic rate per leaf dry mass and plant relative growth rate. A clear understanding of leaf vein traits provides a new perspective on plant function independently of the LES and can enhance the ability to explain and predict whole plant performance under dynamic conditions, with applications towards breeding improved crop varieties.

Coordinated evolution of leaf and stem economics in tropical dry forest trees.

With data from 15 species in eight families of tropical dry forest trees, we provide evidence of coordination between the stem and leaf economic spectra. Species with low-density, flexible, breakable, hydraulically efficient but cavitationally vulnerable wood shed their leaves rapidly in response to drought and had low leaf mass per area and dry mass content. In contrast, species with the opposite xylem syndrome shed their costlier but more drought-resistant leaves late in the dry season. Our results explain variation in the timing of leaf shedding in tropical dry forests: selection eliminates combinations such as low-productivity leaves atop highly vulnerable xylem or water-greedy leaves supplied by xylem of low conductive efficiency. Across biomes, rather than a fundamental trade-off underlying a single axis of trait covariation, the relationship between leaf and stem economics is likely to occupy a wide space in which multiple combinations are possible.