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      Biomechanics and functional morphology of a climbing monocot

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          Abstract

          Climbing monocots can develop into large bodied plants despite being confined by primary growth. In our study on Flagellaria indica we measured surprisingly high stem biomechanical properties (in bending and torsion) and we show that the lack of secondary growth is overcome by a combination of tissue maturation processes and attachment mode. This leads to higher densities of mechanically relevant tissues in the periphery of the stem and to the transition from self-supporting to climbing growth. The development of specialised attachment structures has probably underpinned the evolution of numerous other large bodied climbing monocot taxa.

          Abstract

          Plants with a climbing growth habit possess unique biomechanical properties arising from adaptations to changing loading conditions connected with close attachment to mechanical supports. In monocot climbers, mechanical adaptation is restricted by the absence of a bifacial vascular cambium. Flagellaria indica was used to investigate the mechanical properties and adaptations of a monocot climber that, uniquely, attaches to the surrounding vegetation via leaf tendrils. Biomechanical methods such as three-point bending and torsion tests were used together with anatomical studies on tissue development, modification and distribution. In general, the torsional modulus was lower than the bending modulus; hence, torsional stiffness was less than flexural stiffness. Basal parts of mature stems showed the greatest stiffness while that of more apical stem segments levelled off. Mechanical properties were modulated via tissue maturation processes mainly affecting the peripheral region of the stem. Peripheral vascular bundles showed a reduction in the amount of conducting tissue while the proportion and density of the bundle sheath increased. Furthermore, adjacent bundle sheaths merged resulting in a dense ring of fibrous tissue. Although F. indica lacks secondary cambial growth, the climbing habit is facilitated by a complex interaction of tissue maturation and attachment.

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          Most cited references17

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          Chemical imaging of poplar wood cell walls by confocal Raman microscopy.

          Confocal Raman microscopy was used to illustrate changes of molecular composition in secondary plant cell wall tissues of poplar (Populus nigra x Populus deltoids) wood. Two-dimensional spectral maps were acquired and chemical images calculated by integrating the intensity of characteristic spectral bands. This enabled direct visualization of the spatial variation of the lignin content without any chemical treatment or staining of the cell wall. A small (0.5 microm) lignified border toward the lumen was observed in the gelatinous layer of poplar tension wood. The variable orientation of the cellulose was also characterized, leading to visualization of the S1 layer with dimensions smaller than 0.5 mum. Scanning Raman microscopy was thus shown to be a powerful, nondestructive tool for imaging changes in molecular cell wall organization with high spatial resolution.
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            Moving with climbing plants from Charles Darwin's time into the 21st century.

            We provide an overview of research on climbing plants from Charles Darwin to the present day. Following Darwin's interests, this review will focus on functional perspectives including attachment mechanisms and stem structure and function. We draw attention to a number of unsolved problems inviting future research. These include the mechanism for establishment of the twining habit, a quantitative description following the development of a tissue element through space and time, the chemistry of sticky exudates, the microstructure of xylem and the capacity for water storage, the vulnerability to embolism, and the mechanism for embolism repair. In conclusion we cite evidence that, in response to increasing CO(2) concentration, anthropic perturbation and/ or increasing forest fragmentation, lianas are increasing relative to tree species. In the 21st century, we are returning to the multiscale, multidisciplinary approach taken by Darwin to understand natural history.
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              Global diversification of a tropical plant growth form: environmental correlates and historical contingencies in climbing palms

              Tropical rain forests (TRF) are the most diverse terrestrial biome on Earth, but the diversification dynamics of their constituent growth forms remain largely unexplored. Climbing plants contribute significantly to species diversity and ecosystem processes in TRF. We investigate the broad-scale patterns and drivers of species richness as well as the diversification history of climbing and non-climbing palms (Arecaceae). We quantify to what extent macroecological diversity patterns are related to contemporary climate, forest canopy height, and paleoclimatic changes. We test whether diversification rates are higher for climbing than non-climbing palms and estimate the origin of the climbing habit. Climbers account for 22% of global palm species diversity, mostly concentrated in Southeast Asia. Global variation in climbing palm species richness can be partly explained by past and present-day climate and rain forest canopy height, but regional differences in residual species richness after accounting for current and past differences in environment suggest a strong role of historical contingencies in climbing palm diversification. Climbing palms show a higher net diversification rate than non-climbers. Diversification analyses of palms detected a diversification rate increase along the branches leading to the most species-rich clade of climbers. Ancestral character reconstructions revealed that the climbing habit originated between early Eocene and Miocene. These results imply that changes from non-climbing to climbing habits may have played an important role in palm diversification, resulting in the origin of one fifth of all palm species. We suggest that, in addition to current climate and paleoclimatic changes after the late Neogene, present-day diversity of climbing palms can be explained by morpho-anatomical innovations, the biogeographic history of Southeast Asia, and/or ecological opportunities due to the diversification of high-stature dipterocarps in Asian TRFs.
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                Author and article information

                Journal
                AoB Plants
                AoB Plants
                aobpla
                aobpla
                AoB Plants
                Oxford University Press
                2041-2851
                2016
                27 January 2016
                : 8
                : plw005
                Affiliations
                [1 ]Institut für Botanik, Technische Universität Dresden , D-01062 Dresden, Germany
                [2 ]Institut für Spezielle Botanik, Johannes Gutenberg—Universität Mainz , D-55112 Mainz, Germany
                [3 ]Present address: Plant Biomechanics Group Freiburg, University of Freiburg, Schänzlestrasse 1, D-79104 Freiburg im Breisgau, Germany
                Author notes
                [* ]Corresponding author's e-mail address: linnea.hesse@ 123456biologie.uni-freiburg.de

                Associate Editor: Michael B. Jackson

                Article
                plw005
                10.1093/aobpla/plw005
                4789545
                26819259
                b2ed425f-2cb8-49c1-ac5c-404ff5b43fb4
                Published by Oxford University Press on behalf of the Annals of Botany Company.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 24 April 2015
                : 14 January 2016
                Page count
                Pages: 16
                Categories
                1003
                1029
                Research Articles

                Plant science & Botany
                biomechanics,climbing plants,flagellaria indica,functional morphology,monocotyledons,structural bending modulus,structural modulus of torsion,three-point bending,twist-to-bend ratio

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