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Showing papers in "American Journal of Botany in 2006"


Journal ArticleDOI
TL;DR: The evolution of vessels in angiosperm wood may have required early angiosperms to survive a phase of mechanic and hydraulic instability, and the greater size of the multicellular vessel is facilitated by fibers that strengthen heteroxylous wood.
Abstract: The wide size range of conifer tracheids and angiosperm vessels has important consequences for function. In both conduit types, bigger is better for conducting efficiency. The gain in efficiency with size is maximized by the control of conduit shape, which balances end-wall and lumen resistances. Although vessels are an order of magnitude longer than tracheids of the same diameter, they are not necessarily more efficient because they lack the low end-wall resistance of tracheids with torus-margo pits. Instead, vessels gain conducting efficiency over tracheids by achieving wider maximum diameters. End-walls contributed 56-64% to total xylem resistance in both conduit types, indicating that length limits conducting efficiency. Tracheid dimensions may be more limited by unicellularity and the need to supply strength to homoxylous wood than by the need to protect against cavitation. In contrast, the greater size of the multicellular vessel is facilitated by fibers that strengthen heteroxylous wood. Vessel dimensions may be most limited by the need to restrict intervessel pitting and cavitation by air-seeding. Stressful habitats that promote narrow vessels should favor coexistence of conifers and angiosperms. The evolution of vessels in angiosperm wood may have required early angiosperms to survive a phase of mechanic and hydraulic instability.

550 citations


Journal ArticleDOI
TL;DR: To estimate the evolutionary history of the mustard family, 113 species were sampled, representing 101 of the roughly 350 genera and 17 of the 19 tribes of the family, for the chloroplast gene ndhF and the Shimodaira-Hasegawa test (S-H test) was used to examine the utility of trichome branching patterns for describing monophyletic groups in the ndhf phylogeny.
Abstract: To estimate the evolutionary history of the mustard family (Brassicaceae or Cruciferae), we sampled 113 species, representing 101 of the roughly 350 genera and 17 of the 19 tribes of the family, for the chloroplast gene ndhF. The included accessions increase the number of genera sampled over previous phylogenetic studies by four-fold. Using parsimony, likelihood, and Bayesian methods, we reconstructed the phylogeny of the gene and used the Shimodaira-Hasegawa test (S-H test) to compare the phylogenetic results with the most recent tribal classification for the family. The resultant phylogeny allowed a critical assessment of variations in fruit morphology and seed anatomy, upon which the current classification is based. We also used the S-H test to examine the utility of trichome branching patterns for describing monophyletic groups in the ndhF phylogeny. Our phylogenetic results indicate that 97 of 114 ingroup accessions fall into one of 21 strongly supported clades. Some of these clades can themselves be grouped into strongly to moderately supported monophyletic groups. One of these lineages is a novel grouping overlooked in previous phylogenetic studies. Results comparing 30 different scenarios of evolution by the S-H test indicate that five of 12 tribes represented by two or more genera in the study are clearly polyphyletic, although a few tribes are not sampled well enough to establish para- or polyphyly. In addition, branched trichomes likely evolved independently several times in the Brassicaceae, although malpighiaceous and stellate trichomes may each have a single origin.

399 citations


Journal ArticleDOI
TL;DR: The current state of knowledge of a mechanosensory network in plants is reviewed, and two mechanoreceptor models are considered: a plasmodesmata-based cytoskeleton-plasma membrane-cell wall (CPMCW) network vs. stretch-activated ion channels.
Abstract: The perception of mechanical stimuli in the environment is crucial to the survival of all living organisms. Recent advances have led to the proposal of a plant-specific mechanosensory network within plant cells that is similar to the previously described network in animal systems. This sensory network is the basis for a unifying hypothesis, which may account for the perception of numerous mechanical signals including gravitropic, thigmomorphic, thigmotropic, self-loading, growth strains, turgor pressure, xylem pressure potential, and sound. The current state of our knowledge of a mechanosensory network in plants is reviewed, and two mechanoreceptor models are considered: a plasmodesmata-based cytoskeleton-plasma membrane-cell wall (CPMCW) network vs. stretch-activated ion channels. Post-mechanosensory physiological responses to mechanical stresses are also reviewed, and future research directions in the area of mechanoperception and response are recommended.

308 citations


Journal ArticleDOI
TL;DR: The significance of cell wall architecture for allometric growth can be demonstrated by disturbing the oriented deposition of wall polymers with microtubule-interfering drugs such as colchicine.
Abstract: Growth of turgid cells, defined as an irreversible increase in cell volume and surface area, can be regarded as a physical process governed by the mechanical properties of the cell wall and the osmotic properties of the protoplast. Irreversible cell expansion is produced by creating a driving force for water uptake by decreasing the turgor through stress relaxation in the cell wall. This mechano-hydraulic process thus depends on and can be controlled by the mechanical properties of the wall, which in turn are subject to modification by wall loosening and wall stiffening reactions. The biochemical mechanisms of these changes in mechanical wall properties and their regulation by internal signals (e.g., hormones) or external signals (e.g., light, drought stress) are at present incompletely understood and subject to intensive research. These signals act on walls that have the properties of composite materials in which the molecular structure and spatial organization of polymers rather than the distribution of mechanical stresses dictate the allometry of cell and organ growth and thus cell and organ shape. The significance of cell wall architecture for allometric growth can be demonstrated by disturbing the oriented deposition of wall polymers with microtubule-interfering drugs such as colchicine. Elongating organs (e.g., cylindrical stems or coleoptiles) composed of different tissues with different mechanical properties exhibit longitudinal tissue tensions resulting in the transfer of wall stress from inner to peripheral cell layers that adopt control over organ growth. For physically analyzing the growth process leading to seed germination, the same mechanical and hydraulic parameters as in normal growth are principally appropriate. However, for covering the influences of the tissues that restrain embryo expansion (seed coat, endosperm), an additional force and a water permeability term must be considered.

297 citations


Journal ArticleDOI
TL;DR: Plants can show considerable phenotypic plasticity in mechanical traits, allowing adjustment to changing environments across a range of spatial and temporal scales, but it is not always clear whether a mechanical property is adaptive or a consequence of the physiology associated with stress.
Abstract: Fundamental plant traits such as support, anchorage, and protection against environmental stress depend substantially on biomechanical design. The costs, subsequent trade-offs, and effects on plant performance of mechanical traits are not well understood, but it appears that many of these traits have evolved in response to abiotic and biotic mechanical forces and resource deficits. The relationships between environmental stresses and mechanical traits can be specific and direct, as in responses to strong winds, with structural reinforcement related to plant survival. Some traits such as leaf toughness might provide protection from multiple forms of stress. In both cases, the adaptive value of mechanical traits may vary between habitats, so is best considered in the context of the broader growth environment, not just of the proximate stress. Plants can also show considerable phenotypic plasticity in mechanical traits, allowing adjustment to changing environments across a range of spatial and temporal scales. However, it is not always clear whether a mechanical property is adaptive or a consequence of the physiology associated with stress. Mechanical traits do not only affect plant survival; evidence suggests they have downstream effects on ecosystem organization and functioning (e.g., diversity, trophic relationships, and productivity), but these remain poorly explored.

274 citations


Journal ArticleDOI
TL;DR: A new dynamic model of a tree is described, incorporating the dynamic structural properties of the trunk and branches, and it is indicated that sway is not a harmonic, but is very complex due to the dynamic interaction of branches.
Abstract: Tree stability in windstorms and tree failure are important issues in urban areas where there can be risks of damage to people and property and in forests where wind damage causes economic loss. Current methods of managing trees, including pruning and assessment of mechanical strength, are mainly based on visual assessment or the experience of people such as trained arborists. Only limited data are available to assess tree strength and stability in winds, and most recent methods have used a static approach to estimate loads. Recent research on the measurement of dynamic wind loads and the effect on tree stability is giving a better understanding of how different trees cope with winds. Dynamic loads have been measured on trees with different canopy shapes and branch structures including a palm (Washingtonia robusta), a slender Italian cypress (Cupressus sempervirens) and trees with many branches and broad canopies including hoop pine (Araucaria cunninghamii) and two species of eucalypt (Eucalyptus grandis, E. teretecornus). Results indicate that sway is not a harmonic, but is very complex due to the dynamic interaction of branches. A new dynamic model of a tree is described, incorporating the dynamic structural properties of the trunk and branches. The branch mass contributes a dynamic damping, termed mass damping, which acts to reduce dangerous harmonic sway motion of the trunk and so minimizes loads and increases the mechanical stability of the tree. The results from 12 months of monitoring sway motion and wind loading forces are presented and discussed.

262 citations


Journal ArticleDOI
TL;DR: The cox1 gene has the potential to be used for DNA barcoding of red algae, although a good taxonomic foundation coupled with extensive sampling of taxa is essential for the development of an effective identification system.
Abstract: The red algae, a remarkably diverse group of organisms, are difficult to identify using morphology alone. Following the proposal to use the mitochondrial cytochrome c oxidase subunit I (cox1) for DNA barcoding animals, we assessed the use of this gene in the identification of red algae using 48 samples plus 31 sequences obtained from GenBank. The data set spanned six orders of red algae: the Bangiales, Ceramiales, Corallinales, Gigartinales, Gracilariales and Rhodymeniales. The results indicated that species could be discriminated. Intraspecific variation was between 0 and 4 bp over 539 bp analyzed except in Mastocarpus stellatus (0-14 bp) and Gracilaria gracilis (0-11 bp). Cryptic diversity was found in Bangia fuscopurpurea, Corallina officinalis, G. gracilis, M. stellatus, Porphyra leucosticta and P. umbilicalis. Interspecific variation across all taxa was between 28 and 148 bp, except for G. gracilis and M. stellatus. A comparison of cox1 with the plastid Rubisco spacer for Porphyra species revealed that it was a more sensitive marker in revealing incipient speciation and cryptic diversity. The cox1 gene has the potential to be used for DNA barcoding of red algae, although a good taxonomic foundation coupled with extensive sampling of taxa is essential for the development of an effective identification system.

232 citations


Journal ArticleDOI
TL;DR: Evidence is found that publication bias, experimental design, and the response variable chosen all influence the magnitude of pollen limitation, and suggestions for reducing the bias introduced by methodology in pollen supplementation experiments are offered.
Abstract: Our understanding of pollen limitation depends on a realistic view of its magnitude. Previous reviews of pollen supplementation experiments concluded that a majority of plant species suffers from pollen limitation and that its magnitude is high. Here, we perform a meta-analysis and find evidence that publication bias, experimental design, and the response variable chosen all influence the magnitude of pollen limitation. Fail-safe numbers indicate that publication bias exists for some measures of pollen limitation; significant results are more likely to be published and therefore available for review. Moreover, experiments conducted on only a fraction of a plant’s flowers and reproductive episodes report ;8-fold higher effect sizes than those on all flowers produced over the entire lifetime, likely because resource reallocation among flowers and across years contributes to estimates of pollen limitation. Studies measuring percentage fruit set report higher values of pollen limitation than those measuring other response variables, such as seeds per fruit, perhaps because many plant species will not produce fruits unless adequate pollen receipt occurs to fertilize most ovules. We offer suggestions for reducing the bias introduced by methodology in pollen supplementation experiments and discuss our results in the context of optimality theory.

208 citations


Journal ArticleDOI
Ingo Burgert1
TL;DR: The micromechanical approaches reviewed here are not exhaustive, but they do provide a reasonably comprehensive overview of the methodology with which the general mechanisms underlying the functionality of plant micro- and nanostructure can be explored without destroying the entire cell wall.
Abstract: Plants are hierarchically organized in a way that their macroscopic properties emerge from their micro- and nanostructural level. Hence, micromechanical investigations, which focus on the mechanical design of plant cell walls, are well suited for elucidating the details of the relationship between plant form and function. However, due to the complex nature of primary and secondary cell walls, micromechanical tests on the entire structure cannot provide exact values for polymer properties but must be targeted at the general mechanisms of cell wall deformation and polymer interaction. The success of micromechanical examinations depends on well-considered specimen selection and/or sample pretreatment as well as appropriate experimental setups. Making use of structural differences by taking advantage of the natural variability in plant tissue and cell structure, adaptation strategies can be analyzed at the micro- and nanoscale. Targeted genetic and enzymatic treatments can be utilized to specifically modify individual polymers without degrading the structural integrity of the cell wall. The mechanical properties of such artificial systems reveal the functional roles of individual polymers for a better understanding of the mechanical interactions within the cell wall assembly. In terms of testing methodology, in situ methods that combine micromechanical testing with structural and chemical analyses are particularly well suited for the study of the basic structure-property relationships in plant design. The micromechanical approaches reviewed here are not exhaustive, but they do provide a reasonably comprehensive overview of the methodology with which the general mechanisms underlying the functionality of plant micro- and nanostructure can be explored without destroying the entire cell wall.

198 citations


Journal ArticleDOI
TL;DR: The robust phylogenetic hypothesis inferred from the PHYA data provides a much better context in which to evaluate the evolution of parasitism within Orobanchaceae, the largest of the parasitic angiosperm families.
Abstract: Partial sequences of the nuclear gene encoding the photoreceptor phytochrome A (PHYA) are used to reconstruct relationships within Orobanchaceae, the largest of the parasitic angiosperm families. The monophyly of Orobanchaceae, including nonphotosynthetic holoparasites, hemiparasites, and nonparasitic Lindenbergia is strongly supported. Phytochrome A data resolve six well-supported lineages that contain all of the sampled genera except Brandisia, which is sister to the major radiation of hemiparasites. In contrast to previous plastid and ITS trees, relationships among these major clades also are generally well supported. Thus, the robust phylogenetic hypothesis inferred from the PHYA data provides a much better context in which to evaluate the evolution of parasitism within the group. Ninety-eight species of Orobanchaceae, representing 43 genera, are included and Brandisia, Bungea, Cymbaria, Esterhazya, Nesogenes, Phtheirospermum, Radamaea, Siphonostegia, and Xylocalyx are confirmed as members of Orobanchaceae. The earliest diverging lineage of hemiparasites is identified for the first time; it contains Bungea, Cymbaria, Monochasma, Siphonostegia, and the monotypic Schwalbea, which is federally endangered. This basal clade is marked by the presence of two novel introns. A second, apparently independent gain of one of these introns marks a clade of largely European taxa. There is significant rate heterogeneity among PHYA sequences, and the presence of multiple PHYA in some taxa is consistent with observed ploidy levels.

189 citations


Journal ArticleDOI
TL;DR: Dated photographs of plants in flower represent a new resource to extend the range of species and localities addressed in global-warming research and show changes in flowering times that closely match independent data on the same species in the same locations.
Abstract: Global warming is affecting natural systems across the world. Of the biological responses to warming, changes in the timing of phenological events such as flowering are among the most sensitive. Despite the recognized importance of phenological changes, the limited number of long-term records of phenological events has restricted research on the topic in most areas of the world. In a previous study in Boston (American Journal of Botany 91: 1260-1264), we used herbarium specimens and one season of field observations to show that plants flowered earlier as the climate warmed over the past 100 yr. In our new study, we found that two extra years of data did not strengthen the explanatory power of the analysis. Analysis of herbarium specimens without any field data yielded results similar to analyses that included field observations. In addition, we found that photographs of cultivated and wild plants in Massachusetts, data similar to that contained in herbarium specimens, show changes in flowering times that closely match independent data on the same species in the same locations. Dated photographs of plants in flower represent a new resource to extend the range of species and localities addressed in global-warming research.

Journal ArticleDOI
TL;DR: The aim of this paper is to review the recent work done related to tree-pulling and wind tunnel experiments and mechanistic modeling approaches to increase the understanding of the mechanical stability of trees under static loading.
Abstract: Wind affects the structure and functioning of a forest ecosystem continuously and may cause significant economic loss in managed forests by reducing the yield of recoverable timber, increasing the cost of unscheduled thinning and clear-cuttings, and creating problems in forestry planning. Furthermore, broken and uprooted trees within the forest are subject to insect attack and may provide a suitable breeding substrate, endangering the remaining trees. Therefore, an improved understanding of the processes behind the occurrence of wind-induced damage is of interest to many forest ecologists, but may also help managers of forest resources to make appropriate management decisions related to risk management. Using fundamental physics, empirical experiments, and mechanistic model-based approaches in interaction, we can study the susceptibility of tree stands to wind damage as affected by the wind and site and tree/stand characteristics and management. Such studies are not possible based on statistical approaches alone, which are not able to define the causal links between tree parameters and susceptibility to wind damage. The aim of this paper is to review the recent work done related to tree-pulling and wind tunnel experiments and mechanistic modeling approaches to increase our understanding of the mechanical stability of trees under static loading.

Journal ArticleDOI
TL;DR: Trait expression influenced pollination more so than robbing and pollen receipt was lower in plants that expressed higher levels of leaf gelsemine in two sites, implying that traits associated with pollination and herbivore resistance may not be independent.
Abstract: Plants interact simultaneously with a diversity of visitors, including herbivores and pollinators. Correlations among traits associated with herbivory and pollination may constrain the degree to which plants can evolve in response to any one interactor. Using the distylous plant, Gelsemium sempervirens , we tested the hypothesis that traits typically associated with pollination (distyly) and herbivore resistance (secondary compounds) were phenotypically correlated and examined how these traits influenced plant interactions with floral visitors. The flowers of G. sempervirens are visited by pollinators and a nectar robber, and the leaves and flowers express gelsemine, an alkaloid that is deterrent and sometimes toxic to visitors. Using an observational approach across five populations, we found the thrum floral morph (short-styled) expressed more leaf gelsemine than the pin morph (long-styled). Leaf gelsemine concentrations were positively correlated with flower gelsemine; however, there were no correlations between gelsemine and other floral morphological traits. Trait expression influenced pollination more so than robbing. Thrums received two times less pollen than pins. Moreover, across both morphs, pollen receipt was lower in plants that expressed higher levels of leaf gelsemine in two sites. These results imply that traits associated with pollination and herbivore resistance may not be independent.

Journal ArticleDOI
TL;DR: Trait plasticities were uncorrelated and independent of the magnitude of the canopy gradient in irradiance or height and of the species' light requirements for regeneration, resulting in bounded plasticity that improves canopy performance.
Abstract: Intracanopy plasticity in tree leaf form is a major determinant of whole-plant function and potentially of forest understory ecology. However, there exists little systematic information for the full extent of intracanopy plasticity, whether it is linked with height and exposure, or its variation across species. For arboretum-grown trees of six temperate deciduous species averaging 13-18 m in height, we quantified intracanopy plasticity for 11 leaf traits across three canopy locations (basal-interior, basal-exterior, and top). Plasticity was pronounced across the canopy, and maximum likelihood analyses indicated that plasticity was primarily linked with irradiance, regardless of height. Intracanopy plasticity (the quotient of values for top and basal-interior leaves) was often similar across species and statistically indistinguishable across species for several key traits. At canopy tops, the area of individual leaves was on average 0.5-0.6 times that at basal-interior, stomatal density 1.1-1.5 times higher, sapwood cross-sectional area up to 1.7 times higher, and leaf mass per area 1.5-2.2 times higher; guard cell and stomatal pore lengths were invariant across the canopy. Species differed in intracanopy plasticity for the mass of individual leaves, leaf margin dissection, ratio of leaf to sapwood areas, and stomatal pore area per leaf area; plasticity quotients ranged only up to ≈2. Across the six species, trait plasticities were uncorrelated and independent of the magnitude of the canopy gradient in irradiance or height and of the species' light requirements for regeneration. This convergence across species indicates general optimization or constraints in development, resulting in a bounded plasticity that improves canopy performance.

Journal ArticleDOI
TL;DR: It is shown that this phenomenon is most likely triggered by irregular droughts based on 10 years of observations, and changes in the ENSO cycle resulting from global warming, may have serious ramifications for forest regeneration in this region.
Abstract: General flowering is a community-wide masting phenomenon, which is thus far documented only in aseasonal tropical forests in Asia. Although the canopy and emergent layers of forests in this region are dominated by species of a single family, Dipterocarpaceae, general flowering involves various plant groups. Studying proximate factors and estimating the flowering patterns of the past and future may aid our understanding of the ecological significance and evolutionary factors behind this phenomenon. Here we show that this phenomenon is most likely triggered by irregular droughts based on 10 years of

Journal ArticleDOI
TL;DR: Sharp prickles and/or stellate hairs evolved more than once in Solanum, and floral heterandry originated multiple times within the ‘‘spiny solanums’’ waxy.
Abstract: Species of Solanum subgenus Leptostemonum comprise almost one third of the genus and are distributed worldwide. Members of this group are defined by their sharp epidermal prickles; thus, they are commonly referred to as the ‘‘spiny solanums.’’ This subgenus includes a number of economically important species such as the Old World eggplants, as well as locally cultivated New World species such as the naranjilla and cocona. Given the size and importance of this group we have examined phylogenetic relationships across subgenus Leptostemonum, including a large sampling of species from previously defined species groups within the subgenus. Evolutionary relationships were inferred using DNA sequence data from two nuclear regions (ITS and the granule-bound starch synthase gene [GBSSI or waxy]) and one chloroplast spacer region (trnS-trnG). Results suggest that Solanum subgenus Leptostemonum is monophyletic when the S. wendlandii and S. nemorense species groups are excluded. We have defined 10 clades within subgenus Leptostemonum, some of which correspond to previously circumscribed species groups or sections. Most of the Old World species of subgenus Leptostemonumbelong to a single species-rich clade. Sharp prickles and/or stellate hairs evolved more than once in Solanum, and floral heterandry originated multiple times within the ‘‘spiny solanums.’’ waxy.

Journal ArticleDOI
TL;DR: This work analyzes and explains why spruce is the preferred choice for soundboards, why tropical species are favored for xylophone bars and woodwind instruments, why violinists still prefer pernambuco over other species as a bow material, and why hornbeam and birch are used in piano actions.
Abstract: The unique mechanical and acoustical properties of wood and its aesthetic appeal still make it the material of choice for musical instruments and the interior of concert halls. Worldwide, several hundred wood species are available for making wind, string, or percussion instruments. Over generations, first by trial and error and more recently by scientific approach, the most appropriate species were found for each instrument and application. Using material property charts on which acoustic properties such as the speed of sound, the characteristic impedance, the sound radiation coefficient, and the loss coefficient are plotted against one another for woods. We analyze and explain why spruce is the preferred choice for soundboards, why tropical species are favored for xylophone bars and woodwind instruments, why violinists still prefer pernambuco over other species as a bow material, and why hornbeam and birch are used in piano actions.

Journal ArticleDOI
TL;DR: The consistency between morphological and molecular data suggests that the current phylogeny provides a solid framework for a formal revision of the generic-level classification and for addressing other aspects of the biology of Bignonieae.
Abstract: The tribe Bignonieae (Bignoniaceae) is a large and morphologically diverse clade of neotropical lianas. Despite being a conspicuous component of the neotropical flora, the systematics of the tribe has remained uncertain due to confusing patterns of morphological variation within the group. Chloroplast (ndhF) and nuclear (PepC) DNA sequences were used here to reconstruct the phylogeny of Bignonieae. Individual analyses of ndhF and PepC were highly similar to one another, yet localized differences in the placement of six species suggests some conflict between data sets. Combined analyses result in trees that are consistent with those from the individual analyses and provide greater support for the suggested relationships. This phylogeny provides important new insights into the systematics of the tribe. It identifies 21 strongly supported species groups, eight of which broadly correspond to currently recognized genera. In addition, each of these 21 species groups is supported by morphological synapomorphies. The consistency between morphological and molecular data suggests that the current phylogeny provides a solid framework for a formal revision of the generic-level classification and for addressing other aspects of the biology of Bignonieae.

Journal ArticleDOI
TL;DR: In this paper, the authors examined the spectral reflectance of Mesoamerican trees for three applications: (1) to compare the magnitude of within and between-species variability in leaf reflectance, (2) to determine the potential for species identification, and (3) to test the strength of relationships between leaf traits (chlorophyll content, mesophyll attributes, thickness) and leaf spectral reflectances.
Abstract: Leaf traits and physiological performance govern the amount of light reflected from leaves at visible and infrared wavebands. Information on leaf optical properties of tropical trees is scarce. Here, we examine leaf reflectance of Mesoamerican trees for three applications: (1) to compare the magnitude of within- and between-species variability in leaf reflectance, (2) to determine the potential for species identification based on leaf reflectance, and (3) to test the strength of relationships between leaf traits (chlorophyll content, mesophyll attributes, thickness) and leaf spectral reflectance. Within species, shape and amplitude differences between spectra were compared within single leaves, between leaves of a single tree, and between trees. We also investigated the variation in a species' leaf reflectance across sites and seasons. Using forward feature selection and pattern recognition tools, species classification within a single site and season was successful, while classification between sites or seasons was not. The implications of variability in leaf spectral reflectance were considered in light of potential tree crown classifications from remote airborne or satellite-borne sensors. Species classification is an emerging field with broad applications to tropical biologists and ecologists, including tree demographic studies and habitat diversity assessments.

Journal ArticleDOI
TL;DR: The pollination biology of a group of 15 orchids that share a recognizable syndrome of floral features that includes yellow-green coloration, oil secretion, pungent scent, shallow flowers, and a September peak in flowering is examined, suggesting that pollinators play an important role in mediating selection on floral traits.
Abstract: The long-standing notion of pollination syndromes, which postulates that plants form recognizable groups according to pollinator type, has been challenged recently on the basis of apparent widespread generalization in pollination systems. As a test of the pollination syndrome concept, I examined the pollination biology of a group of 15 orchids that share a recognizable syndrome of floral features that includes yellow-green coloration, oil secretion, pungent scent, shallow flowers, and a September peak in flowering. The orchids occur in sympatry in the Cape Floral Region of South Africa. According to the pollination syndrome concept, the similar floral features of this group indicate a shared pollinator. To test this prediction, I observed pollinators on Pterygodium alatum, P. caffrum, P. catholicum, P. volucris, Corycium orobanchoides, and Disperis bolusiana subsp. bolusiana. They shared a single species of pollinator, the oil-collecting bee, Rediviva peringueyi. Female bees collected oil from the lip appendage using modified front tarsi. The orchids reduce interspecific reproductive interference through differences in pollinarium length or the use of mutually exclusive pollinarium attachment sites on the body of the bee. The results are contrary to the expectation of generalization in pollination systems and suggest that pollinators play an important role in mediating selection on floral traits.

Journal ArticleDOI
TL;DR: Protection against air-seeding in conifer tracheids appears to be uncoupled from conduit size and conducting efficiency, in contrast to recent work on angiosperm vessels.
Abstract: Plant xylem must balance efficient delivery of water to the canopy against protection from air entry into the conduits via air-seeding. We investigated the relationship between tracheid allometry, end wall pitting, safety from air-seeding, and the hydraulic efficiency of conifer wood in order to better understand the trade-offs between effective transport and protection against air entry. Root and stem wood were sampled from conifers belonging to the Pinaceae, Cupressaceae, Podocarpaceae, and Araucariaceae. Hydraulic resistivity of tracheids decreased with increasing tracheid diameter and width, with 64 ± 4% residing in the end wall pitting regardless of tracheid size or phylogenetic affinity. This end-wall percentage was consistent with a near-optimal scaling between tracheid diameter and length that minimized flow resistance for a given tracheid length. There was no evidence that tracheid size and hydraulic efficiency were constrained by the role of the pits in protecting against cavitation by air-seeding. An increase in pit area resistance with safety from cavitation was observed only for species of the northern hemisphere (Pinaceae and Cupressaceae), but this variable was independent of tracheid size, and the increase in pit resistance did not significantly influence tracheid resistance. In contrast to recent work on angiosperm vessels, protection against air-seeding in conifer tracheids appears to be uncoupled from conduit size and conducting efficiency.

Journal ArticleDOI
TL;DR: This study illustrates how ecophysiological differences can shift the competitive advantage from one species to another along a stress gradient as land use patterns change on the two continents.
Abstract: An invasive variety of Phragmites australis (Poaceae, common reed), the M haplotype, has been implicated in the spread of this species into North American salt marshes that are normally dominated by the salt marsh grass Spartina alterniflora (Poaceae, smooth cordgrass). In some European marshes, on the other hand, Spartina spp. derived from S. alterniflora have spread into brackish P. australis marshes. In both cases, the non-native grass is thought to degrade the habitat value of the marsh for wildlife, and it is important to understand the physiological processes that lead to these species replacements. We compared the growth, salt tolerance, and osmotic adjustment of M haplotype P. australis and S. alterniflora along a salinity gradient in greenhouse experiments. Spartina alterniflora produced new biomass up to 0.6 M NaCl, whereas P. australis did not grow well above 0.2 M NaCl. The greater salt tolerance of S. alterniflora compared with P. australis was due to its ability to use Na(+) for osmotic adjustment in the shoots. On the other hand, at low salinities P. australis produced more shoots per gram of rhizome tissue than did S. alterniflora. This study illustrates how ecophysiological differences can shift the competitive advantage from one species to another along a stress gradient. Phragmites australis is spreading into North American coastal marshes that are experiencing reduced salinities, while Spartina spp. are spreading into northern European brackish marshes that are experiencing increased salinities as land use patterns change on the two continents.

Journal ArticleDOI
TL;DR: Phylogenetic analysis of the combined data provided a fairly robust phylogeny for Vitaceae, which shows complex multiple intercontinental relationships within the northern hemisphere and between northern and southern hemispheres.
Abstract: Seventy-nine species representing 12 genera of Vitaceae were sequenced for the trnL-F spacer, 37 of which were subsequently sequenced for the atpB-rbcL spacer and the rps16 intron. Phylogenetic analysis of the combined data provided a fairly robust phylogeny for Vitaceae. Cayratia, Tetrastigma, and Cyphostemma form a clade. Cyphostemma and Tetrastigma are each monophyletic, and Cayratia may be paraphyletic. Ampelopsis is paraphyletic with the African Rhoicissus and the South American Cissus striata nested within it. The pinnately leaved Ampelopsis form a subclade, and the simple and palmately leaved Ameplopsis constitutes another with both subclades containing Asian and American species. Species of Cissus from Asia and Central America are monophyletic, but the South American C. striata does not group with other Cissus species. The Asian endemic Nothocissus and Pterisanthes form a clade with Asian Ampelocissus, and A. javalensis from Central America is sister to this clade. Vitis is monophyletic and forms a larger clade with Ampelocissus, Pterisanthes, and Nothocissus. The eastern Asian and North American disjunct Parthenocissus forms a clade with Yua austro-orientalis, a species of a small newly recognized genus from China to eastern Himalaya. Vitaceae show complex multiple intercontinental relationships within the northern hemisphere and between northern and southern hemispheres.

Journal ArticleDOI
TL;DR: This work investigates the phylogeny of the Caryophyllaceae by means of analyzing plastid and nuclear sequence data with parsimony and Bayesian methods and describes a mode of tracing a stable phylogenetic signal in ITS sequences.
Abstract: Caryophyllaceae is a principally holarctic family including around 2200 species often classified into the three subfamilies Alsinoideae, Caryophylloideae, and Paronychioideae. Complex and possibly homoplasious morphological characters within the family make taxa difficult to delimit and diagnose. To explore part of the morphological evolution within the family, we investigated the phylogeny of the Caryophyllaceae by means of analyzing plastid and nuclear sequence data with parsimony and Bayesian methods. We describe a mode of tracing a stable phylogenetic signal in ITS sequences, and a significant common signal is shared with the plastid data. Parsimony and Bayesian analyses yield some differences in tree resolution. None of the subfamilies appear monophyletic, but the monophyly of the Caryophylloideae is not contradicted. Alsinoideae are paraphyletic, with Arenaria subg. Eremogone and Minuartia subg. Spergella more closely related to the Caryophylloideae. There is strong support for the inclusion of Spergula-Spergularia in an Alsinoideae-Caryophylloideae clade. Putative synapomorphies for these groupings are twice as many stamens as number of sepals and a caryophyllad-type of embryogeny. Paronychioideae form a basal grade, where tribe Corrigioleae are sister to the rest of the family. Free styles and capsules with simple teeth are possibly plesiomorphic for the family.

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TL;DR: Structural and functional constraints leading to the trade-off of efficiency against safety of water transport were demonstrated at the individual pit level, with a positive correlation between pit membrane resistance on an area basis and the pressure differential required to cause membrane stretching, a characteristic that is essential for pit aspiration.
Abstract: The air-seeding hypothesis predicts that xylem embolism resistance is linked directly to bordered pit functioning. We tested this prediction in trunks, roots, and branches at different vertical and radial locations in young and old trees of Pseudotsuga menziesii. Dimensions of bordered pits were measured from light and scanning electron micrographs, and physiological data were from published values. Consistent with observations, calculations showed that earlywood tracheids were more resistant to embolism than latewood tracheids, mainly from earlywood having stretchier pit membranes that can distend and cover the pit aperture. Air seeding that occurs in earlywood appears to happen through gaps between the torus edge and pit border, as shown by the similar calculated pressures required to stretch the membrane over the pit aperture and to cause embolism. Although bordered pit functioning was correlated with tracheid hydraulic diameter, pit pore size and above all pit aperture constrained conductivity the most. From roots to branches and from the trunk base to higher on the trunk, hydraulic resistance of the earlywood pit membrane increased significantly because of a decrease in the size of the pit aperture and size and number of margo pores. Moreover, overall wood conductivity decreased, in part due to lower pit conductivity and a decrease in size and frequency of pits. Structural and functional constraints leading to the trade-off of efficiency against safety of water transport were also demonstrated at the individual pit level, with a positive correlation between pit membrane resistance on an area basis and the pressure differential required to cause membrane stretching, a characteristic that is essential for pit aspiration.

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TL;DR: The significance of inspiration from nature for technical textiles and for fibrous composite materials is demonstrated by examples of already existing technical solutions that either parallel biology or are indeed inspired by biological models.
Abstract: The significance of inspiration from nature for technical textiles and for fibrous composite materials is demonstrated by examples of already existing technical solutions that either parallel biology or are indeed inspired by biological models. The two different basic types of biomimetic approaches are briefly presented and discussed for the "technical plant stem." The technical plant stem is a biomimetic product inspired by a variety of structural and functional properties found in different plants. The most important botanical templates are the stems of the giant reed (Arundo donax, Poaceae) and of the Dutch rush (Equisetum hyemale, Equisetaceae). After analysis of the structural and mechanical properties of these plants, the physical principles have been deduced and abstracted and finally transferred to technical applications. Modern computer-controlled fabrication methods for producing technical textiles and for structuring the embedding matrix of compound materials render unique possibilities for transferring the complex structures found in plants, which often are optimized on several hierarchical levels, into technical applications. This process is detailed for the technical plant stem, a biomimetic, lightweight, fibrous composite material based on technical textiles with optimized mechanical properties and a gradient structure.

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TL;DR: Progress in the understanding of plant posture and mechanical acclimation will require new biomechanical models of plant architectural development, as well as new types of kinematic design.
Abstract: Self-supporting plant stems are slender, erect structures that remain standing while growing in highly variable mechanical environments. Such ability is not merely related to an adapted mechanical design in terms of material-specific stiffness and stem tapering. As many terrestrial standing animals do, plant stems regulate posture through active and coordinated control of motor systems and acclimate their skeletal growth to prevailing loads. This analogy probably results from mechanical challenges on standing organisms in an aerial environment with low buoyancy and high turbulence. But the continuous growth of plants submits them to a greater challenge. In response to these challenges, land plants implemented mixed skeletal and motor functions in the same anatomical elements. There are two types of kinematic design: (1) plants with localized active movement (arthrophytes) and (2) plants with continuously distributed active movements (contortionists). The control of these active supporting systems involves gravi- and mechanoperception, but little is known about their coordination at the whole plant level. This more active view of the control of plant growth and form has been insufficiently considered in the modeling of plant architecture. Progress in our understanding of plant posture and mechanical acclimation will require new biomechanical models of plant architectural development.

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TL;DR: The results suggest that flowers were ancestrally monos asymmetric with seven fertile stamens and three adaxial staminodes, switched to asymmetry later, and reverted to monosymmetry in clade VII, and suggest these nectaries may represent a key innovation in plant defense strategies that enabled Senna to undergo large-scale diversification.
Abstract: Senna (Leguminosae) is a large, widespread genus that includes species with enantiostylous, asymmetric flowers and species with extrafloral nectaries. Clarification of phylogenetic relationships within Senna based on parsimony analyses of three chloroplast regions (rpS16, rpL16, and matK) provides new insights on the evolution of floral symmetry and extrafloral nectaries. Our results support the monophyly of only one (Psilorhegma) of the six currently recognized sections, while Chamaefistula, Peiranisia, and Senna are paraphyletic, and monotypic Astroites and Paradictyon are nested within two of the seven major clades identified by our molecular phylogeny. Two clades (I, VII) include only species with monosymmetric flowers, while the remaining clades (II-VI) contain species with asymmetric, enantiostylous flowers, in which either the gynoecium alone or, in addition, corolla and androecium variously contribute to the asymmetry. Our results further suggest that flowers were ancestrally monosymmetric with seven fertile stamens and three adaxial staminodes, switched to asymmetry later, and reverted to monosymmetry in clade VII. Fertility of all 10 stamens is a derived state, characterizing the Psilorhegma subclade. Extrafloral nectaries evolved once and constitute a synapomorphy for clades IV-VII ("EFN clade"). These nectaries may represent a key innovation in plant defense strategies that enabled Senna to undergo large-scale diversification.

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TL;DR: Little evidence is found that environmental gradients drive patterns of fern species richness on this spatial scale, and mean annual rainfall and temperature, light intensities in the canopy and at ground level, and the mid-domain effect are examined.
Abstract: To better understand changes in the distribution and diversity within plant functional types along an elevational gradient and the potential mechanisms driving such changes, we studied species richness of ferns at six elevations along a forested elevational gradient in Costa Rica, from La Selva Biological Station at 30 m a.s.l. up the slopes of Volcan Barva to 2960 m a.s.l. Among the samples from all the sites combined, we found 264 species from 60 genera. Sixty-nine species were terrestrial, 113 were canopy epiphytes, and 121 were low-trunk epiphytes. Only one species occupied both canopy and terrestrial habitats at any of the study sites. Overlap of canopy and low-trunk species composition was relatively low (18%), and lower still was the overlap of terrestrial and low-trunk species (12%). Total species richness peaked at the 1000-m site for canopy and low-trunk epiphytic species. In contrast, the richness of terrestrial species rose to a mid-elevation maximum and remained relatively constant at higher elevations. In an effort to explain elevational patterns of species richness, we examined mean annual rainfall and temperature, light intensities in the canopy and at ground level, and the mid-domain effect. Of the explanatory factors examined, the mid-domain effect accounted for most of the elevational pattern. We found little evidence that environmental gradients drive patterns of fern species richness on this spatial scale.

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TL;DR: The results are consistent with a scenario in which black walnut recolonized its current range from a single glacial refugium, and where subsequent genetic effects associated with deforestation and habitat fragmentation have been mitigated by high levels of pollen flow.
Abstract: Broad-scale studies of genetic structure and diversity are indicative of the recent evolutionary history of a species and are relevant to conservation efforts. We have estimated current levels of genetic diversity and population structure for black walnut (Juglans nigra L.), a highly valuable timber species, in the central hardwood region of the United States. Black walnut trees from 43 populations across this region were genotyped at 12 highly polymorphic microsatellite loci. Genetic diversity was high and populations only slightly deviated from Hardy-Weinberg proportions (FIS= 0.017). Considering the scale of our sampling, the species was remarkably genetically homogenous: FST was quite low (0.017), and in a Bayesian analysis the optimal higher-order partition was into a single group comprised of all 43 populations. Although black walnut is predominantly a bottomland species, very little genetic variance was partitioned among broad-scale hydrologic regions (FPT= 0.002). However, a weak, but statistically significant pattern of isolation by distance was detected. The results are consistent with a scenario in which black walnut recolonized its current range from a single glacial refugium, and where subsequent genetic effects associated with deforestation and habitat fragmentation have been mitigated by high levels of pollen flow. Nuclear microsatellites alone may be insufficient to identify hotspots for black walnut conservation.