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allenai

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scirepeval

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30666535

10.1111/J.1365-2435.2010.01698.X

Rethinking the value of high wood density

Summary 1. Current thinking holds that wood density mediates a tradeoff between strength and economy of construction, with higher wood density providing higher strength but at higher cost. 2. Yet the further away wood fibres are from the central axis of the trunk, the more they increase the strength of the trunk; thus, a fat trunk of low-density wood can achieve greater strength at lower construction cost than a thin trunk of high-density wood. 3. What then are the countervailing advantages of high wood density? 4. We hypothesize that high wood density is associated with lower maintenance costs due to lower trunk surface area, as surface area correlates with maintenance respiration. 5. This advantage would be particularly important to long-lived trees and could in part explain why they tend to have high wood density. 6. High wood density has also been associated with lower risk of trunk breakage, xylem implosion and pathogen invasion, but we argue that these relationships are not causal and instead reflect correlated selection on other traits of value to long-lived trees. 7. This revaluation of the costs and benefits of high wood density has important implications for understanding tree life-history evolution, functional diversity, forest carbon stocks and the impacts of global change.

Functional Ecology

31503318

10.1002/BIES.20597

The echinoderm collagen fibril: a hero in the connective tissue research of the 1990s

Collagen fibrils are some of the most‐abundant and important extracellular structures in our bodies, yet we are unsure of their shape and size. This is largely due to an inherent difficulty in isolating them from their surrounding tissues. Echinoderms have collagenous tissues that are similar to ours in many ways, yet they can be manipulated to easily relinquish their collagen fibrils, providing an excellent opportunity to study native fibrillar structure. In the early 1990s, they were found to defy the commonly accepted fibrillar model of the time in that they were much shorter, they were shaped like double‐ended spindles, and their centers exhibited a reversal in molecular polarity. Realization of these features helped to reform the questions that were being asked about vertebrate fibrils, shifting the focus toward shape and size. Since then, researchers working with both groups (echinoderms and vertebrates) have worked together to find the structure of native fibrils. This information will be fundamental in understanding what holds collagenous tissues together at the fibrillar level, and could have important implications for people with Ehlers‐Danlos syndrome. BioEssays 29:645–653, 2007. © 2007 Wiley Periodicals, Inc.

BioEssays

45157909

10.1111/PCE.12325

Implications of the mesophyll conductance to CO2 for photosynthesis and water-use efficiency during long-term water stress and recovery in two contrasting Eucalyptus species.

Water stress (WS) slows growth and photosynthesis (A(n)), but most knowledge comes from short-time studies that do not account for longer term acclimation processes that are especially relevant in tree species. Using two Eucalyptus species that contrast in drought tolerance, we induced moderate and severe water deficits by withholding water until stomatal conductance (g(sw)) decreased to two pre-defined values for 24 d, WS was maintained at the target g(sw) for 29 d and then plants were re-watered. Additionally, we developed new equations to simulate the effect on mesophyll conductance (g(m)) of accounting for the resistance to refixation of CO(2). The diffusive limitations to CO(2), dominated by the stomata, were the most important constraints to A(n). Full recovery of A(n) was reached after re-watering, characterized by quick recovery of gm and even higher biochemical capacity, in contrast to the slower recovery of g(sw). The acclimation to long-term WS led to decreased mesophyll and biochemical limitations, in contrast to studies in which stress was imposed more rapidly. Finally, we provide evidence that higher gm under WS contributes to higher intrinsic water-use efficiency (iWUE) and reduces the leaf oxidative stress, highlighting the importance of gm as a target for breeding/genetic engineering.

Plant Cell and Environment

107333470

10.1098/RSPB.2003.2637

Evidence for an elastic projection mechanism in the chameleon tongue

To capture prey, chameleons ballistically project their tongues as far as 1.5 body lengths with accelerations of up to 500 m s⊃–2. At the core of a chameleon's tongue is a cylindrical tongue skeleton surrounded by the accelerator muscle. Previously, the cylindrical accelerator muscle was assumed to power tongue projection directly during the actual fast projection of the tongue. However, high–speed recordings of Chamaeleo melleri and C. pardalis reveal that peak powers of 3000 W kg⊃–1 are necessary to generate the observed accelerations, which exceed the accelerator muscle's capacity by at least five– to 10–fold. Extrinsic structures might power projection via the tongue skeleton. High–speed fluoroscopy suggests that they contribute less than 10% of the required peak instantaneous power. Thus, the projection power must be generated predominantly within the tongue, and an energy–storage–and–release mechanism must be at work. The key structure in the projection mechanism is probably a cylindrical connective–tissue layer, which surrounds the entoglossal process and was previously suggested to act as lubricating tissue. This tissue layer comprises at least 10 sheaths that envelop the entoglossal process. The outer portion connects anteriorly to the accelerator muscle and the inner portion to the retractor structures. The sheaths contain helical arrays of collagen fibres. Prior to projection, the sheaths are longitudinally loaded by the combined radial contraction and hydrostatic lengthening of the accelerator muscle, at an estimated mean power of 144 W kg⊃–1 in C. melleri. Tongue projection is triggered as the accelerator muscle and the loaded portions of the sheaths start to slide over the tip of the entoglossal process. The springs relax radially while pushing off the rounded tip of the entoglossal process, making the elastic energy stored in the helical fibres available for a simultaneous forward acceleration of the tongue pad, accelerator muscle and retractor structures. The energy release continues as the multilayered spring slides over the tip of the smooth and lubricated entoglossal process. This sliding–spring theory predicts that the sheaths deliver most of the instantaneous power required for tongue projection. The release power of the sliding tubular springs exceeds the work rate of the accelerator muscle by at least a factor of 10 because the elastic–energy release occurs much faster than the loading process. Thus, we have identified a unique catapult mechanism that is very different from standard engineering designs. Our morphological and kinematic observations, as well as the available literature data, are consistent with the proposed mechanism of tongue projection, although experimental tests of the sheath strain and the lubrication of the entoglossal process are currently beyond our technical scope.

Proceedings of The Royal Society B: Biological Sciences

96091028

10.1371/JOURNAL.PONE.0000395

Magnetoreception System in Honeybees (Apis mellifera)

Honeybees (Apis mellifera) undergo iron biomineralization, providing the basis for magnetoreception. We showed earlier the presence of superparamagnetic magnetite in iron granules formed in honeybees, and subscribed to the notion that external magnetic fields may cause expansion or contraction of the superparamagnetic particles in an orientation-specific manner, relaying the signal via cytoskeleton (Hsu and Li 1994). In this study, we established a size-density purification procedure, with which quantitative amount of iron granules was obtained from honey bee trophocytes and characterized; the density of iron granules was determined to be 1.25 g/cm3. While we confirmed the presence of superparamagnetic magnetite in the iron granules, we observed changes in the size of the magnetic granules in the trophycytes upon applying additional magnetic field to the cells. A concomitant release of calcium ion was observed by confocal microscope. This size fluctuation triggered the increase of intracellular Ca+2 , which was inhibited by colchicines and latrunculin B, known to be blockers for microtubule and microfilament syntheses, respectively. The associated cytoskeleton may thus relay the magnetosignal, initiating a neural response. A model for the mechanism of magnetoreception in honeybees is proposed, which may be applicable to most, if not all, magnetotactic organisms.

PLOS ONE

343461

10.1080/09687680701446965

The structural basis of water permeation and proton exclusion in aquaporins (Review)

Aquaporins (AQPs) represent a ubiquitous class of integral membrane proteins that play critical roles in cellular osmoregulations in microbes, plants and mammals. AQPs primarily function as water-conducting channels, whereas members of a sub-class of AQPs, termed aquaglyceroporins, are permeable to small neutral solutes such as glycerol. While AQPs facilitate transmembrane permeation of water and/or small neutral solutes, they preclude the conduction of protons. Consequently, openings of AQP channels allow rapid water diffusion down an osmotic gradient without dissipating electrochemical potentials. Molecular structures of AQPs portray unique features that define the two central functions of AQP channels: effective water permeation and strict proton exclusion. This review describes AQP structures known to date and discusses the mechanisms underlying water permeation, proton exclusion and water permeability regulation.

Molecular Membrane Biology

86026645

10.1890/0012-9658(1997)078[1946:PANEOO]2.0.CO;2

POSITIVE AND NEGATIVE EFFECTS OF ORGANISMS AS PHYSICAL ECOSYSTEM ENGINEERS

Physical ecosystem engineers are organisms that directly or indirectly control the availability of resources to other organisms by causing physical state changes in biotic or abiotic materials. Physical ecosystem engineering by organisms is the physical modification, maintenance, or creation of habitats. Ecological effects of engineers on many other species occur in virtually all ecosystems because the physical state changes directly create nonfood resources such as living space, directly control abiotic resources, and indirectly modulate abiotic forces that, in turn, affect resource use by other organisms. Trophic interactions and resource competition do not constitute engineering. Engineering can have significant or trivial effects on other species, may involve the physical structure of an organism (like a tree) or structures made by an organism (like a beaver dam), and can, but does not invariably, have feedback effects on the engineer. We argue that engineering has both negative and positive effects on species richness and abundances at small scales, but the net effects are probably positive at larger scales encompassing engineered and nonengineered environments in ecological and evolutionary space and time. Models of the population dynamics of engineers suggest that the engineer/habitat equilibrium is often, but not always, locally stable and may show long-term cycles, with potential ramifications for community and ecosystem stability. As yet, data adequate to parameterize such a model do not exist for any engineer species. Because engineers control flows of energy and materials but do not have to participate in these flows, energy, mass, and stoichiometry do not appear to be useful in predicting which engineers will have big effects. Empirical observations suggest some potential generalizations about which species will be important engineers in which ecosystems. We point out some of the obvious, and not so obvious, ways in which engineering and trophic relations interact, and we call for greater research on physical ecosystem engineers, their impacts, and their interface with trophic relations.

Ecology

31876174

10.1002/ANIE.200803189

Blue luminescence of ripening bananas.

As revealed in the last two decades, chlorophyll breakdown in senescent leaves appears to occur by a largely common and well-controlled catabolic path, which rapidly furnishes non-fluorescent, colorless chlorophyll catabolites (NCCs) as “final” products.[1,2] Recently NCCs detected in ripe apples and pears were found to be the same as those in degreened leaves, suggesting chlorophyll catabolism in leaf senescence and fruit ripening to be similar (Scheme 1).[3,4]

Angewandte Chemie

90418337

10.1111/JAV.01294

Contributions of feather microstructure to eider down insulation properties

Insulation is an essential component of nest structure that helps provide incubation requirements for birds. Many species of waterfowl breed in high latitudes where rapid heat loss can necessitate a high energetic input from parents and use down feathers to line their nests. Common eider Somateria mollissima nest down has exceptional insulating properties but the microstructural mechanisms behind the feather properties have not been thoroughly examined. Here, we hypothesized that insulating properties of nest down are correlated to down feather (plumule) microstructure. We tested the thermal efficiency (fill power) and cohesion of plumules from nests of two Icelandic colonies of wild common eiders and compared them to properties of plumules of wild greylag goose Anser anser. We then used electron microscopy to examine the morphological basis of feather insulating properties. We found that greylag goose down has higher fill power (i.e. traps more air) but much lower cohesion (i.e. less prone to stick together) compared to common eider down. These differences were related to interspecific variation in feather microstructure. Down cohesion increased with the number of barbule microstructures (prongs) that create strong points of contact among feathers. Eider down feathers also had longer barbules than greylag goose down feathers, likely increasing their air-trapping capacity. Feather properties of these two species might reflect the demands of their contrasting evolutionary history. In greylag goose, a temperate, terrestrial species, plumule microstructure may optimize heat trapping. In common eiders, a diving duck that nests in arctic and subarctic waters, plumule structure may have evolved to maximize cohesion over thermal insulation, which would both reduce buoyancy during their foraging dives and enable nest down to withstand strong arctic winds.

Journal of Avian Biology

710555

10.1016/J.JSB.2011.07.008

²⁶Mg labeling of the sea urchin regenerating spine: Insights into echinoderm biomineralization process.

This paper reports the results of the first dynamic labeling experiment with regenerating spines of sea urchins Paracentrotus lividus using the stable isotope ²⁶Mg and NanoSIMS high-resolution isotopic imaging, which provide a direct information about the growth process. Growing spines were labeled twice (for 72 and 24 h, respectively) by increasing the abundance of ²⁶Mg in seawater. The incorporation of ²⁶Mg into the growing spines was subsequently imaged with the NanoSIMS ion microprobe. Stereom trabeculae initially grow as conical micro-spines, which form within less than 1 day. These micro-spines fuse together by lateral outgrowths and form a thin, open meshwork (inner stereom), which is subsequently reinforced by addition of layered thickening deposits (outer stereom). The (longitudinal) growth rate of the inner stereom is ca. 125 μm/day. A single (ca. 1 μm) thickening layer in the stereom trabeculae is deposited during 24h. The thickening process is contemporaneous with the formation micro-spines and involves both longitudinal trabeculae and transverse bridges to a similar degree. Furthermore, the skeleton-forming cells remain active in the previously formed open stereom for at least 10 days, and do not migrate upwards until the end of the thickening process. The experimental capability presented here provides a new way to obtain detailed information about the skeleton formation of a multitude of marine, calcite producing organisms.

Journal of Structural Biology

6160984

10.1128/AEM.65.4.1477-1482.1999

Microbiology of the Oil Fly, Helaeomyia petrolei

ABSTRACT Helaeomyia petrolei larvae isolated from the asphalt seeps of Rancho La Brea in Los Angeles, Calif., were examined for microbial gut contents. Standard counts on Luria-Bertani, MacConkey, and blood agar plates indicated ca. 2 × 105 heterotrophic bacteria per larva. The culturable bacteria represented 15 to 20% of the total population as determined by acridine orange staining. The gut itself contained large amounts of the oil, had no observable ceca, and maintained a slightly acidic pH of 6.3 to 6.5. Despite the ingestion of large amounts of potentially toxic asphalt by the larvae, their guts sustained the growth of 100 to 1,000 times more bacteria than did free oil. All of the bacteria isolated were nonsporeformers and gram negative. Fourteen isolates were chosen based on representative colony morphologies and were identified by using the Enterotube II and API 20E systems and fatty acid analysis. Of the 14 isolates, 9 were identified as Providencia rettgeriand 3 were likely Acinetobacter isolates. No evidence was found that the isolates grew on or derived nutrients from the asphalt itself or that they played an essential role in insect development. Regardless, any bacteria found in the oil fly larval gut are likely to exhibit pronounced solvent tolerance and may be a future source of industrially useful, solvent-tolerant enzymes.

Applied and Environmental Microbiology

29866867

10.1007/BF00173949

Dispersal-generated sexual selection in a beetle-riding pseudoscorpion

SummaryAfter several generations within a decaying tree (Ficus spp.), populations of the pseudoscorpion Cordylochernes scorpioides disperse by climbing under the elytra of harlequin beetles (Acrocinus longimanus) eclosing from the tree. Because the beetles then fly to newly-decaying Ficus for mating and oviposition, they act as effective dispersal agents. Field experiments and observations indicate that this dispersal mode has been exploited by males who compete to remain on a beetle as a strategic site for inseminating females dispersing on it and on other beetles. Whereas beetles just eclosed from old trees carried large, female-biased groups of dispersing pseudoscorpions, beetles captured after their maiden flight generally carried a single large male. Multivariate morphometric analyses indicated that these beetle-riding males were much larger than individuals randomly sampled from trees, with fighting traits exhibiting the greatest potential for selection (highest “phoretic” differentials). In females, sexual receptivity was significantly higher at the beginning of dispersal than at the end, suggesting that mating occurs on beetles. Field experiments confirmed that on-beetle insemination does take place and that small males are displaced from the “subelytral space” by larger rivals. By contrast, laboratory experiments suggest that large male size may not confer high mating success under the low-density conditions which characterize populations within trees. Thus, in addition to indicating a novel role for dispersal in the evolution of exaggerated male traits, this study suggests that oscillating sexual selection may be important in the maintenance of the extreme phenotypic variation exhibited by males of this species.

Behavioral Ecology and Sociobiology

124704265

10.1063/1.4942474

Direct numerical simulation of sharkskin denticles in turbulent channel flow

The hydrodynamic function of sharkskin has been under investigation for the past 30 years. Current literature conflicts on whether sharkskin is able to reduce skin friction similar to riblets. To contribute insights toward reconciling these conflicting views, direct numerical simulations are carried out to obtain detailed flow fields around realistic denticles. A sharp interface immersed boundary method is employed to simulate two arrangements of actual sharkskin denticles (from Isurus oxyrinchus) in a turbulent boundary layer at Reτ ≈ 180. For comparison, turbulent flow over drag-reducing scalloped riblets is also simulated with similar flow conditions and with the same numerical method. Although the denticles resemble riblets, both sharkskin arrangements increase total drag by 44%-50%, while the riblets reduce drag by 5%. Analysis of the simulated flow fields shows that the turbulent flow around denticles is highly three-dimensional and separated, with 25% of the total drag being form drag. The complex three-dimensional shape of the denticles gives rise to a mean flow dominated by strong secondary flows in sharp contrast with the mean flow generated by riblets, which is largely two-dimensional. The so resulting three-dimensionality of sharkskin flows leads to an increase in the magnitude of the turbulent statistics near the denticles, which further contributes to increasing the total drag. The simulations also show that, at least for the simulated arrangements, sharkskin, in sharp contrast with drag-reducing riblets, is unable to isolate high shear stress near denticle ridges causing a significant portion of the denticle surface to be exposed to high mean shear.

Physics of Fluids

481163

10.1242/JEB.033159

Spatial vision in the purple sea urchin Strongylocentrotus purpuratus (Echinoidea)

SUMMARY Recent evidence that echinoids of the genus Echinometra have moderate visual acuity that appears to be mediated by their spines screening off-axis light suggests that the urchin Strongylocentrotus purpuratus, with its higher spine density, may have even more acute spatial vision. We analyzed the movements of 39 specimens of S. purpuratus after they were placed in the center of a featureless tank containing a round, black target that had an angular diameter of 6.5 deg. or 10 deg. (solid angles of 0.01 sr and 0.024 sr, respectively). An average orientation vector for each urchin was determined by testing the animal four times, with the target placed successively at bearings of 0 deg., 90 deg., 180 deg. and 270 deg. (relative to magnetic east). The urchins showed no significant unimodal or axial orientation relative to any non-target feature of the environment or relative to the changing position of the 6.5 deg. target. However, the urchins were strongly axially oriented relative to the changing position of the 10 deg. target (mean axis from –1 to 179 deg.; 95% confidence interval ± 12 deg.; P<0.001, Moore's non-parametric Hotelling's test), with 10 of the 20 urchins tested against that target choosing an average bearing within 10 deg. of either the target center or its opposite direction (two would be expected by chance). In addition, the average length of the 20 target-normalized bearings for the 10 deg. target (each the vector sum of the bearings for the four trials) were far higher than would be expected by chance (P<10−10; Monte Carlo simulation), showing that each urchin, whether it moved towards or away from the target, did so with high consistency. These results strongly suggest that S. purpuratus detected the 10 deg. target, responding either by approaching it or fleeing it. Given that the urchins did not appear to respond to the 6.5 deg. target, it is likely that the 10 deg. target was close to the minimum detectable size for this species. Interestingly, measurements of the spine density of the regions of the test that faced horizontally predicted a similar visual resolution (8.3±0.5 deg. for the interambulacrum and 11±0.54 deg. for the ambulacrum). The function of this relatively low, but functional, acuity – on par with that of the chambered Nautilus and the horseshoe crab – is unclear but, given the bimodal response, is likely to be related to both shelter seeking and predator avoidance.

The Journal of Experimental Biology

84210172

10.4039/ENT96264-1

Temporal Patterns of Emergence in Aquatic Insects

In aquatic insects, emergence (ecdysis to the adult or subimaginal stage) varies widely in temporal pattern. The comparative study of this pattern is feasible and informative in orders such as Plecoptera, Ephemeroptera, Odonata, Diptera and Trichoptera in which all members of a population pass through the water-surface when emerging. Methods by which emergence rate can be measured are discussed. Four basic temporal patterns of emergence exist. Emergence may be (1) continuous with irregular fluctuations in rate; (2) rhythmic, with a lunar period; (3) sporadic, occurring at irregular intervals of a few days; or (4) seasonal. Examples of each of these patterns are given, and reference is made to the proximate and ultimate environmental factors which may be maintaining the patterns observed. Diurnal rhythms of emergence are excluded from consideration. When emergence is restricted seasonally in temperate latitudes, the degree of its synchronization within the emergence period varies widely but is usually constant and typical for a given species. This has provided the basis for an ecological classification of British Odonata, the validity of which is examined in the light of recent research.

Canadian Entomologist

7576513

10.1093/ICB/ICN069

How kelp produce blade shapes suited to different flow regimes: A new wrinkle.

Many species of macroalgae have flat, strap-like blades in habitats exposed to rapidly flowing water, but have wide, ruffled "undulate" blades at protected sites. We used the giant bull kelp, Nereocystis luetkeana, to investigate how these ecomorphological differences are produced. The undulate blades of N. luetkeana from sites with low flow remain spread out and flutter erratically in moving water, thereby not only enhancing interception of light, but also increasing drag. In contrast, strap-like blades of kelp from habitats with rapid flow collapse into streamlined bundles and flutter at low amplitude in flowing water, thus reducing both drag and interception of light. Transplant experiments in the field revealed that shape of the blade in N. luetkeana is a plastic trait. Laboratory experiments in which growing blades from different sites were subjected to tensile forces that mimicked the hydrodynamic drag experienced by blades in different flow regimes showed that change in shape is induced by mechanical stress. During growth experiments in the field and laboratory, we mapped the spatial distribution of growth in both undulate and strap-like blades to determine how these different morphologies were produced. The highest growth rates occur near the proximal ends of N. luetkeana blades of both morphologies, but the rates of transverse growth of narrow, strap-like blades are lower than those of wide, undulate blades. If rates of longitudinal growth at the edges of a blade exceed the rate of longitudinal growth along the midline of the blade, ruffles along the edges of the blade are produced by elastic buckling. In contrast, flat blades are produced when rates of longitudinal growth are similar across the width of a blade. Because ruffles are the result of elastic buckling, a compliant undulate N. luetkeana blade can easily be pushed into different configurations (e.g., the wavelengths of the ruffles along the edges of the blade can change, and the whole blade can twist into left- and right-handed helicoidal shapes), which may enhance movements of the blade in flowing water that reduce self-shading and increase mass exchange along blade surfaces.

Integrative and Comparative Biology

55617967

10.4081/JLIMNOL.2004.S1.16

Experiences with dormancy in tardigrades

Tardigrades often colonise extreme habitats, in which they survive using both types of dormancy: quiescence and diapause. Together with nematodes and bdelloid rotifers, tardigrades are known to enter quiescence (with several forms of cryptobiosis: anhydrobiosis, cryobiosis, anoxybiosis, osmobiosis) at any stage of their life cycle, from egg to adult. Entering anhydrobiosis, tardigrades contract their body into a so-called tun, loosing most of their free and bound water (>95%), synthesizing cell protectants (e.g., trehalose, glycerol, heat shock proteins) and strongly reducing or suspending their metabolism. Our research on cryptobiosis focused on some ecological and evolutionary aspects. We evaluated: i) the long-term anhydrobiotic survival by comparing quantitative data on recovery from naturally induced desiccation in several species of tardigrades; ii) differences in survival patterns between species and populations by experimentally inducing anhydrobiosis and cryobiosis; iii) phenotypic factors affecting anhydrobiotic survival. As regards diapause, we considered encystment and eggs. Encystment involves at least the synthesis of new cuticular structures. Morphological changes during cyst formation are more complex than those involved in tun formation. We analyzed more in detail encystment processes, comparing a semiterrestrial with a limnic species. Several inter-specific differences have been identified, other than the production of two types of cysts in the semiterrestrial species. Our analysis of life history traits of a laboratory reared strain of a soil tardigrade revealed a particular hatching phenology that involved the production of both subitaneous and resting eggs. The latter need a cue to hatch (dehydration followed by re-hydration). In addition, the evolutionary meaning of dormancy in tardigrades is discussed.

Journal of Limnology

25973068

10.1007/PL00008153

Simulating ozone detoxification in the leaf apoplast through the direct reaction with ascorbate

Abstract. This paper presents a mathematical model which enables the semi-quantification of ozone (O3) detoxification, based upon the direct reaction of the pollutant with ascorbate (ASC) located in the aqueous matrix associated with the cell wall (i.e. the apoplast). The model describes the uptake of ozone into the leaf and its direct reaction with ASC, taking into consideration the regeneration of dehydroascorbic acid in the cytosol, the rate of replenishment of cell wall ASC and the distribution of ASC between sub-cellular compartments – based upon the permeability of biomembranes to the neutral species, ascorbic acid and the pH of various sub-cellular compartments. The importance of various physico-chemical characteristics (e.g. stomatal conductance, mesophyll cell wall thickness and tortuosity, chloroplast volume, apoplast pH, ASC:O3 reaction stoichiometry) in mediating the flux of ozone to the plasmalemma is analysed. Model simulations, supported by experimental observations, suggest that the ASC concentration in the leaf apoplast is high enough to scavenge a significant proportion of the O3 taken up into the leaf interior, under environmentally relevant conditions. However, there is considerable variation between taxa in the potential degree of protection afforded by apoplastic ASC, emphasizing the need for an improved understanding of the reaction chemistry of O3 in the cell wall.

Planta

14794337

10.1007/S00114-011-0853-5

A novel marine silk

The discovery of a novel silk production system in a marine amphipod provides insights into the wider potential of natural silks. The tube-building corophioid amphipod Crassicorophium bonellii produces from its legs fibrous, adhesive underwater threads that combine barnacle cement biology with aspects of spider silk thread extrusion spinning. We characterised the filamentous silk as a mixture of mucopolysaccharides and protein deriving from glands representing two distinct types. The carbohydrate and protein silk secretion is dominated by complex β-sheet structures and a high content of charged amino acid residues. The filamentous secretion product exits the gland through a pore near the tip of the secretory leg after having moved through a duct, which subdivides into several small ductules all terminating in a spindle-shaped chamber. This chamber communicates with the exterior and may be considered the silk reservoir and processing/mixing space, in which the silk is mechanically and potentially chemically altered and becomes fibrous. We assert that further study of this probably independently evolved, marine arthropod silk processing and secretion system can provide not only important insights into the more complex arachnid and insect silks but also into crustacean adhesion cements.

Naturwissenschaften

42898148

10.1126/SCIENCE.1140097

The Role of Wheat Awns in the Seed Dispersal Unit

The dispersal unit of wild wheat bears two pronounced awns that balance the unit as it falls. We discovered that the awns are also able to propel the seeds on and into the ground. The arrangement of cellulose fibrils causes bending of the awns with changes in humidity. Silicified hairs that cover the awns allow propulsion of the unit only in the direction of the seeds. This suggests that the dead tissue is analogous to a motor. Fueled by the daily humidity cycle, the awns induce the motility required for seed dispersal

Science

1736589

10.1098/RSIF.2009.0203

Quorum sensing and social networking in the microbial world

For many years, bacterial cells were considered primarily as selfish individuals, but, in recent years, it has become evident that, far from operating in isolation, they coordinate collective behaviour in response to environmental challenges using sophisticated intercellular communication networks. Cell-to-cell communication between bacteria is mediated by small diffusible signal molecules that trigger changes in gene expression in response to fluctuations in population density. This process, generally referred to as quorum sensing (QS), controls diverse phenotypes in numerous Gram-positive and Gram-negative bacteria. Recent advances have revealed that bacteria are not limited to communication within their own species but are capable of ‘listening in’ and ‘broadcasting to’ unrelated species to intercept messages and coerce cohabitants into behavioural modifications, either for the good of the population or for the benefit of one species over another. It is also evident that QS is not limited to the bacterial kingdom. The study of two-way intercellular signalling networks between bacteria and both uni- and multicellular eukaryotes as well as between eukaryotes is just beginning to unveil a rich diversity of communication pathways.

Journal of the Royal Society Interface

12280258

10.1371/JOURNAL.PBIO.0050092

Protein Oxidation Implicated as the Primary Determinant of Bacterial Radioresistance

In the hierarchy of cellular targets damaged by ionizing radiation (IR), classical models of radiation toxicity place DNA at the top. Yet, many prokaryotes are killed by doses of IR that cause little DNA damage. Here we have probed the nature of Mn-facilitated IR resistance in Deinococcus radiodurans, which together with other extremely IR-resistant bacteria have high intracellular Mn/Fe concentration ratios compared to IR-sensitive bacteria. For in vitro and in vivo irradiation, we demonstrate a mechanistic link between Mn(II) ions and protection of proteins from oxidative modifications that introduce carbonyl groups. Conditions that inhibited Mn accumulation or Mn redox cycling rendered D. radiodurans radiation sensitive and highly susceptible to protein oxidation. X-ray fluorescence microprobe analysis showed that Mn is globally distributed in D. radiodurans, but Fe is sequestered in a region between dividing cells. For a group of phylogenetically diverse IR-resistant and IR-sensitive wild-type bacteria, our findings support the idea that the degree of resistance is determined by the level of oxidative protein damage caused during irradiation. We present the case that protein, rather than DNA, is the principal target of the biological action of IR in sensitive bacteria, and extreme resistance in Mn-accumulating bacteria is based on protein protection.

PLOS Biology

1241783

10.1186/1742-9994-7-23

Fog-basking behaviour and water collection efficiency in Namib Desert Darkling beetles

BackgroundIn the Namib Desert fog represents an alternative water source. This is utilised by Darkling beetles (Tenebrionidae) that employ different strategies for obtaining the fog water. Some dig trenches in the sand, while others use their own bodies as fog collectors assuming a characteristic fog-basking stance. Two beetle species from the genus Onymacris have been observed to fog-bask on the ridges of the sand dunes. These beetles all have smooth elytra surfaces, while another species with elytra covered in bumps is reported to have specialised adaptations facilitating water capture by fog-basking. To resolve if these other beetles also fog-bask, and if an elytra covered in bumps is a more efficient fog water collector than a smooth one, we examined four Namib Desert beetles; the smooth Onymacris unguicularis and O. laeviceps and the bumpy Stenocara gracilipes and Physasterna cribripes. Here we describe the beetles' fog-basking behaviour, the details of their elytra structures, and determine how efficient their dorsal surface areas are at harvesting water from fog.ResultsThe beetles differ greatly in size. The largest P. cribripes has a dorsal surface area that is 1.39, 1.56, and 2.52 times larger than O. unguicularis, O. laeviceps, and S. gracilipes, respectively. In accordance with earlier reports, we found that the second largest O. unguicularis is the only one of the four beetles that assumes the head standing fog-basking behaviour, and that fog is necessary to trigger this behaviour. No differences were seen in the absolute amounts of fog water collected on the dorsal surface areas of the different beetles. However, data corrected according to the sizes of the beetles revealed differences. The better fog water harvesters were S. gracilipes and O. unguicularis while the large P. cribripes was the poorest. Examination of the elytra microstructures showed clear structural differences, but the elytra of all beetles were found to be completely hydrophobic.ConclusionsThe differences in fog water harvesting efficiency by the dorsal surface areas of beetles with very different elytra surface structures were minor. We therefore conclude that the fog-basking behaviour itself is a more important factor than structural adaptations when O. unguicularis collect water from fog.

Frontiers in Zoology

5778081

10.1186/S13750-016-0060-0

How effective are created or restored freshwater wetlands for nitrogen and phosphorus removal? A systematic review

BackgroundEutrophication of aquatic environments is a major environmental problem in large parts of the world. In Europe, EU legislation (the Water Framework Directive and the Marine Strategy Framework Directive), international conventions (OSPAR, HELCOM) and national environmental objectives emphasize the need to reduce the input of nutrients to freshwater and marine environments. A widely used method to achieve this is to allow water to pass through a created or restored wetland. However, the large variation in measured nutrient removal rates in such wetlands calls for a systematic review.MethodsSearches for primary studies were performed in electronic databases and on the internet. One author performed the screening of all retrieved articles at the title and abstract level. To check that the screening was consistent and complied with the agreed inclusion/exclusion criteria, subsets of 100 articles were screened by the other authors. When screening at full-text level the articles were evenly distributed among the authors. Kappa tests were used to evaluate screening consistency. Relevant articles remaining after screening were critically appraised and assigned to three quality categories, from two of which data were extracted. Quantitative synthesis consists of meta-analyses and response surface analyses. Regressions were performed using generalized additive models that can handle nonlinear relationships and interaction effects.ResultsSearches generated 5853 unique records. After screening on relevance and critical appraisal, 93 articles including 203 wetlands were used for data extraction. Most of the wetlands were situated in Europe and North America. The removal rate of both total nitrogen (TN) and total phosphorus (TP) is highly dependent on the loading rate. Significant relationships were also found for annual average air temperature (T) and wetland area (A). Median removal rates of TN and TP were 93 and 1.2 g m−2 year−1, respectively. Removal efficiency for TN was significantly correlated with hydrologic loading rate (HLR) and T, and the median was 37 %, with a 95 % confidence interval of 29–44 %. Removal efficiency for TP was significantly correlated with inlet TP concentration, HLR, T, and A. Median TP removal efficiency was 46 % with a 95 % confidence interval of 37–55 %. Although there are small differences in average values between the two quality categories, the variation is considerably smaller among high quality studies compared to studies with lower quality. This suggests that part of the large variation between studies may be explained by less rigorous study designs.ConclusionsOn average, created and restored wetlands significantly reduce the transport of TN and TP in treated wastewater and urban and agricultural runoff, and may thus be effective in efforts to counteract eutrophication. However, restored wetlands on former farmland were significantly less efficient than other wetlands at TP removal. In addition, wetlands with precipitation-driven HLRs and/or hydrologic pulsing show significantly lower TP removal efficiencies compared to wetlands with controlled HLRs. Loading rate (inlet concentrations × hydraulic loading rates) needs to be carefully estimated as part of the wetland design. More research is needed on the effects of hydrologic pulsing on wetlands. There is also a lack of evidence for long-term (>20 years) performance of wetlands.

Environmental Evidence

7497510

10.1242/JEB.033365

Insect eggs protected from high temperatures by limited homeothermy of plant leaves

SUMMARY Virtually all aspects of insect biology are affected by body temperature, and many taxa have evolved sophisticated temperature-control mechanisms. All insects, however, begin life as eggs and lack the ability to thermoregulate. Eggs laid on leaves experience a thermal environment, and thus a body temperature, that is strongly influenced by the leaves themselves. Because plants can maintain leaf temperatures that differ from ambient, e.g. by evapotranspiration, plant hosts may protect eggs from extreme ambient temperatures. We examined the degree to which leaves buffer ambient thermal variation and whether that buffering benefits leaf-associated insect eggs. In particular, we: (1) measured temperature variation at oviposition sites in the field, (2) manipulated temperatures in the laboratory to determine the effect of different thermal conditions on embryo development time and survival, and (3) tested embryonic metabolic rates over increasing temperatures. Our results show that Datura wrightii leaves buffer Manduca sexta eggs from fatally high ambient temperatures in the southwestern USA. Moreover, small differences in temperature profiles among leaves can cause large variation in egg metabolic rate and development time. Specifically, large leaves were hotter than small leaves during the day, reaching temperatures that are stressfully high for eggs. This study provides the first mechanistic demonstration of how this type of leaf-constructed thermal refuge interacts with egg physiology.

The Journal of Experimental Biology

29684143

10.1007/S11104-006-9033-3

Allelochemicals released by rice roots and residues in soil

A few rice (Oryza sativa L.) varieties or rice straw produce and release allelochemicals into soil in which interfere with the growth of neighboring or successive plants. Allelopathic rice PI312777 and Huagan-1 at their early growth stages released momilactone B, 3-isopropyl-5-acetoxycyclohexene-2-one-1, and 5,7,4′-trihydroxy-3′,5′-dimethoxyflavone into soil at phytotoxic levels, but non-allelopathic rice Huajingxian did not. Both allelopathic and non-allelopathic rice residues released momilactone B and lignin-related phenolic acids (p-hydroxybenzoic, p-coumaric, ferulic, syringic and vanillic acids) into the soil during residue decomposition to inhibit successive plants. The results indicated that allelochemicals involved in rice allelopathy from living and dead plants are substantially different. Interestingly, the concentrations of the allelochemicals released from the allelopathic rice seedlings in soil increased dramatically when they were surrounded with Echinochloa crus-galli. The concentrations of the allelochemicals were over 3-fold higher in the presence of E. crus-galli than in the absence of E. crus-galli. However, the same case did not occur in non-allelopathic Huajingxian seedlings surrounded with E. crus-galli. In addition to allelochemical exudation being promoted by the presence of E. crus-galli, allelopathic rice seedlings also increased allelochemical exudation in response to exudates of germinated E. crus-galli seeds or lepidimoide, an uronic acid derivative exuded from E. crus-galli seeds. These results imply that allelopathic rice seedlings can sense certain allelochemicals released by E. crus-galli into the soil, and respond by increased production of allelochemicals inhibitory to E. crus-galli. This study suggests that rice residues of both allelopathic and non-allelopathic varieties release similar concentrations and types of allelochemicals to inhibit successive plants. In contrast, living rice plants of certain allelopathic varieties appear to be able to detect the presence of interspecific neighbors and respond by increased allelochemicals.

Plant and Soil

4411789

10.1038/35086574

Crossflow filtration in suspension-feeding fishes

Rows of comb-like or tufted gill rakers in the oral cavity of suspension-feeding fishes (for example, herring, anchovies and tilapia) have been thought to serve as (1) non-porous barriers that direct particle-laden water to the sticky oral roof, where particles are retained as water exits from the oral cavity, (2) conventional dead-end filters that sieve particles from water exiting between rakers, or (3) sticky filters that retain particles encountered by a hydrosol filtration mechanism. Here we present data from computational fluid dynamics and video endoscopy in suspension-feeding fish indicating that the rakers of three distantly related species function instead as a crossflow filter. Particles are concentrated inside the oral cavity as filtrate exits between the rakers, but particles are not retained on the rakers. Instead, the high-velocity crossflow along the rakers carries particles away from the raker surfaces and transports the particles towards the oesophagus. This crossflow prevents particles from clogging the gaps between the rakers, and solves the mystery of particle transport from the rakers to the oesophagus.

Nature

7058398

10.1172/JCI64416

Cytokeratins mediate epithelial innate defense through their antimicrobial properties.

Epithelial cells express antimicrobial proteins in response to invading pathogens, although little is known regarding epithelial defense mechanisms during healthy conditions. Here we report that epithelial cytokeratins have innate defense properties because they constitutively produce cytoprotective antimicrobial peptides. Glycine-rich C-terminal fragments derived from human cytokeratin 6A were identified in bactericidal lysate fractions of human corneal epithelial cells. Structural analysis revealed that these keratin-derived antimicrobial peptides (KDAMPs) exhibited coil structures with low α-helical content. Synthetic analogs of these KDAMPS showed rapid bactericidal activity against multiple pathogens and protected epithelial cells against bacterial virulence mechanisms, while a scrambled peptide showed no bactericidal activity. However, the bactericidal activity of a specific KDAMP was somewhat reduced by glycine-alanine substitutions. KDAMP activity involved bacterial binding and permeabilization, but the activity was unaffected by peptide charge or physiological salt concentration. Knockdown of cytokeratin 6A markedly reduced the bactericidal activity of epithelial cell lysates in vitro and increased the susceptibility of murine corneas to bacterial adherence in vivo. These data suggest that epithelial cytokeratins function as endogenous antimicrobial peptides in the host defense against infection and that keratin-derived antimicrobials may serve as effective therapeutic agents.

Journal of Clinical Investigation

2848034

10.1007/S00359-010-0556-Y

Material properties of the skin of the Kenyan sand boa Gongylophis colubrinus (Squamata, Boidae)

On the basis of structural data, it has been previously assumed that the integument of snakes consists of a hard, robust, inflexible outer surface (Oberhäutchen and β-layer) and soft, flexible inner layers (α-layers). The aim of this study was to compare material properties of the outer and inner scale layers of the exuvium of Gongylophis colubrinus, to relate the structure of the snake integument to its mechanical properties. The nanoindentation experiments have demonstrated that the outer scale layers are harder, and have a higher effective elastic modulus than the inner scale layers. The results obtained provide strong evidence about the presence of a gradient in the material properties of the snake integument. The possible functional significance of this gradient is discussed here as a feature minimizing damage to the integument during sliding locomotion on an abrasive surface, such as sand.

Journal of Comparative Physiology A-neuroethology Sensory Neural and Behavioral Physiology

4327277

10.1038/35090573

Calcitic microlenses as part of the photoreceptor system in brittlestars

Photosensitivity in most echinoderms has been attributed to ‘diffuse’ dermal receptors. Here we report that certain single calcite crystals used by brittlestars for skeletal construction are also a component of specialized photosensory organs, conceivably with the function of a compound eye. The analysis of arm ossicles in Ophiocoma showed that in light-sensitive species, the periphery of the labyrinthic calcitic skeleton extends into a regular array of spherical microstructures that have a characteristic double-lens design. These structures are absent in light-indifferent species. Photolithographic experiments in which a photoresist film was illuminated through the lens array showed selective exposure of the photoresist under the lens centres. These results provide experimental evidence that the microlenses are optical elements that guide and focus the light inside the tissue. The estimated focal distance (4–7 µm below the lenses) coincides with the location of nerve bundles—the presumed primary photoreceptors. The lens array is designed to minimize spherical aberration and birefringence and to detect light from a particular direction. The optical performance is further optimized by phototropic chromatophores that regulate the dose of illumination reaching the receptors. These structures represent an example of a multifunctional biomaterial that fulfills both mechanical and optical functions.

Nature

8690966

10.1177/0748730402239676

Photoperiodic Adjustments in Immune Function Protect Siberian Hamsters from Lethal Endotoxemia

Seasonal changes in day length enhance or suppress components of immune function in individuals of several mammalian species. Siberian hamsters (Phodopus sungorus) exhibit multiple changes in neuroendocrine, reproductive, and immune function after exposure to short days. The manner in which these changes are integrated into the host response to pathogens is not well understood. The present experiments tested the hypothesis that short-day changes in immune function alter the pathogenesis of septic shock and survival after challenge with endotoxin. Male and female Siberian hamsters raised in long-day photoperiods were transferred as adults to short days or remained in their natal photoperiod. Six to 8 weeks later, hamsters were injected i.p. with 0, 1, 2.5, 10, 25, or 50 mg/kg bacterial lipopolysaccharide (LPS) (the biologically active constituent of endotoxin), and survival was monitored for 96 h. Short days significantly improved survival of male hamsters treated with 10 or 25 mg/kg LPS and improved survival in females treated with 50 mg/kg LPS. Transfer from long to short days shifted the LD5 0 in males by approximately 90%, from 5.3 to 9.9 mg/kg, and in females from 11.1 to 15.0 mg/kg ([#002B]35%). Long-day females were more resistant than were males to lethal endotoxemia. In vitro production of the proinflammatory cytokine TNFα in response to LPS stimulation was significantly lower in macrophages extracted from short-day relative to long-day hamsters, as were circulating concentrations of TNFα in vivo after i.p. administration of LPS, suggesting that diminished cytokine responses to LPS in short days may mitigate the lethality of endotoxemia. Adaptation to short days induces changes in immune parameters that affect survival in the face of immune challenges.

Journal of Biological Rhythms

25325987

10.1073/PNAS.1000429107

FM echolocating bats shift frequencies to avoid broadcast–echo ambiguity in clutter

Sonar broadcasts are followed by echoes at different delays from objects at different distances. When broadcasts are emitted rapidly in cluttered surroundings, echo streams from successive broadcasts overlap and cause ambiguity in matching echoes to corresponding broadcasts. To identify reactions to ambiguity in clutter, echolocating bats that emit multiple-harmonic FM sounds were trained to fly into a dense, extended array of obstacles (multiple rows of vertically hanging chains) while the sonar sounds the bat emitted were recorded with a miniature radio microphone carried by the bat. Flight paths were reconstructed from thermal-infrared video recordings. Successive rows of chains extended more than 6 m in depth, so each broadcast was followed by a series of echoes from multiple rows of chains that lasted up to 40 ms. Bats emitted sounds in pairs (“strobe groups”) at short (20–40 ms) interpulse intervals (IPIs) alternating with longer IPIs (>50 ms). For many short IPIs, the stream of echoes from the first broadcast was still arriving when the second broadcast was emitted. This overlap caused ambiguity about matching echoes with broadcasts. Bats shifted frequencies of the first sound in each strobe group upward and the second sound downward by 3–6 kHz. When overlap and ambiguity ceased, frequency shifts ceased also. Frequency differences were small compared with the total broadcast band, which was 75–80 kHz wide, but the harmonic structure of echoes enhances the differences in spectrograms. Bats could use time–frequency comparisons of echoes with broadcasts to assign echoes to the corresponding broadcasts and thus avoid ambiguity.

Proceedings of the National Academy of Sciences of the United States of America

10877733

10.1242/JEB.043661

Always on the bright side of life: anti-adhesive properties of insect ommatidia grating

SUMMARY The surface of some insect eyes consists of arrays of cuticular protuberances, which are 50–300 nm in diameter, and are termed corneal nipples or ommatidia gratings. They were widely reported to reduce the reflectance for normally incident light, contributing to camouflage by reducing glare to predators, while furthermore enhancing the intake of light, which is especially important for nocturnal insects. Our preliminary observations suggest a third function: in contrast to the rest of the body, ommatidia of various insects remain clean, even in a heavy contaminated environment. In order to prove such an anti-contamination hypothesis of these structures, we measured the adhesive properties of polymer moulds of insect ommatidia, and compared these data with control surfaces having the same curvature radii but lacking such a nanostructure. A scanning electron microscope (SEM) study and force measurements using an atomic force microscope (AFM) on the eye surfaces of three different insect species, dragonfly Aeshna mixta (Odonata), moth Laothoe populi (Lepidoptera) and fly Volucella pellucens (Diptera), were undertaken. We revealed that adhesion is greatly reduced by corneal grating in L. populi and V. pellucens when compared with their smooth controls. The smooth cornea of A. mixta showed no statistically significant difference to its control. We assume that this anti-adhesive phenomenon is due to a decrease in the real contact area between contaminating particles and the eye's surface. Such a combination of three functions in one nanostructure can be interesting for the development of industrial multifunctional surfaces capable of enhancing light harvesting while reducing light reflection and adhesion.

The Journal of Experimental Biology

2345933

10.1104/PP.105.075523

Contribution of the Alternative Pathway to Respiration during Thermogenesis in Flowers of the Sacred Lotus1

We report results from in vivo measurements, using oxygen isotope discrimination techniques, of fluxes through the alternative and cytochrome respiratory pathways in thermogenic plant tissue, the floral receptacle of the sacred lotus (Nelumbo nucifera). Fluxes through both pathways were measured in thermoregulating flowers undergoing varying degrees of thermogenesis in response to ambient temperature. Significant increases in alternative pathway flux were found in lotus receptacles with temperatures 16°C to 20°C above ambient, but not in those with lesser amounts of heating. Alternative pathway flux in the hottest receptacles was 75% of the total respiratory flux. In contrast, fluxes through the cytochrome pathway did not change significantly during thermogenesis. These data support the hypothesis that increased flux through the alternative pathway is responsible for heating in the lotus and that it is unlikely that uncoupling proteins, which would have produced increased fluxes through the cytochrome pathway, contribute significantly to heating in this tissue. Comparisons of actual flux, with capacity determined using inhibitors, suggested that the alternative pathway was operating at close to maximum capacity in heating tissues of lotus. However, in nonheating tissues the inhibitor data significantly overestimated the alternative pathway flux. This confirms that isotopic measurements are necessary for accurate determination of fluxes through the two pathways.

Plant Physiology

16735242

10.1371/JOURNAL.PONE.0045735

Catapulting Tentacles in a Sticky Carnivorous Plant

Among trapping mechanisms in carnivorous plants, those termed ‘active’ have especially fascinated scientists since Charles Darwin’s early works because trap movements are involved. Fast snap-trapping and suction of prey are two of the most spectacular examples for how these plants actively catch animals, mainly arthropods, for a substantial nutrient supply. We show that Drosera glanduligera, a sundew from southern Australia, features a sophisticated catapult mechanism: Prey animals walking near the edge of the sundew trigger a touch-sensitive snap-tentacle, which swiftly catapults them onto adjacent sticky glue-tentacles; the insects are then slowly drawn within the concave trap leaf by sticky tentacles. This is the first detailed documentation and analysis of such catapult-flypaper traps in action and highlights a unique and surprisingly complex mechanical adaptation to carnivory.

PLOS ONE

9792930

10.1007/S002210100774

Chameleons have independent eye movements but synchronise both eyes during saccadic prey tracking

Abstract. The movements of both eyes and the head were recorded with search coils in unrestrained, freely moving chameleons. As a main result I found that the generation of saccades in the left and the right eye was either independent from each other or was highly correlated according to the behavioural situation. When no prey item was fixated, disconjugate saccades were observed which was in accordance with earlier observations in chameleons. During prey tracking the chameleons switched to a different oculomotor behaviour and pursued the moving prey with synchronous saccades. At higher target velocities, the tracking movement of the head was also saccadic and was synchronised with the two eyes. Binocular coupling affected only the timing of the saccades but not the metrics: the amplitudes of the synchronous saccades were usually different in the two eyes. These observations suggest the existence of two independent premotor neuronal circuits for left and right eye saccadic motor control in the chameleon. Binocular coupling in prey-tracking chameleons is probably achieved by neuronal coupling of these premotor circuits during eye–head coordination. The ability to switch between synchronous and uncoupled saccadic eye movements has not been described for any other vertebrate. This unique ability of the chameleon may help to understand the organisation of the oculomotor system of other vertebrates since evidence for separate left eye and right eye saccade generation and position control has recently also been reported in primates.

Experimental Brain Research

14826000

10.1126/SCIENCE.1141064

Extracellular Proteins Limit the Dispersal of Biogenic Nanoparticles

High–spatial-resolution secondary ion microprobe spectrometry, synchrotron radiation–based Fourier-transform infrared spectroscopy, and polyacrylamide gel analysis demonstrated the intimate association of proteins with spheroidal aggregates of biogenic zinc sulfide nanocrystals, an example of extracellular biomineralization. Experiments involving synthetic zinc sulfide nanoparticles and representative amino acids indicated a driving role for cysteine in rapid nanoparticle aggregation. These findings suggest that microbially derived extracellular proteins can limit the dispersal of nanoparticulate metal-bearing phases, such as the mineral products of bioremediation, that may otherwise be transported away from their source by subsurface fluid flow.

Science

129027636

10.1016/J.GCA.2003.10.039

Physicochemical characterization of the microhabitat of the epibionts associated with Alvinella pompejana, a hydrothermal vent annelid

Abstract Alvinella pompejana is a polychaetous annelid that inhabits narrow tubes along the walls of high-temperature hydrothermal vent chimneys. The worm hosts a rich community of epibiotic bacteria that coats its dorsal surface. Although the worm tube microhabitat is a challenging environment to sample, characterizing the thermal and geochemical regime is important for understanding the ecology of the worm and its bacteria, as the worm spends most of its time inside the tube. We characterized the physicochemical conditions of diffuse hydrothermal flow inside inhabited worm tubes using in situ analysis and wet chemical analysis of discrete water samples. Thermistor probes deployed inside worm tubes measured temperatures ranging from 28.6°C to 84.0°C, while temperatures at tube orifices ranged from 7.5°C to 40.0°C. In situ electrochemical analysis of tube fluids revealed undetectable oxygen (

Geochimica et Cosmochimica Acta

39683005

10.1073/PNAS.96.17.9705

Spray aiming in the bombardier beetle: photographic evidence.

Bombardier beetles, when physically assaulted, eject a hot quinonoid spray from the tip of the abdomen. Photographic evidence is presented demonstrating that the African bombardier beetle, Stenaptinus insignis, can aim its spray in virtually any direction. It can target its individual legs, and even the individual segments of its legs. Moreover, in aiming at a leg, it takes into account the postural orientation of that leg. The beetle is able even to target sites on its back. It is postulated that the ability to aim helps the beetle mainly in defense against ants.

Proceedings of the National Academy of Sciences of the United States of America

9826833

10.1073/PNAS.0900086106

Characterization of an electron conduit between bacteria and the extracellular environment

A number of species of Gram-negative bacteria can use insoluble minerals of Fe(III) and Mn(IV) as extracellular respiratory electron acceptors. In some species of Shewanella, deca-heme electron transfer proteins lie at the extracellular face of the outer membrane (OM), where they can interact with insoluble substrates. To reduce extracellular substrates, these redox proteins must be charged by the inner membrane/periplasmic electron transfer system. Here, we present a spectro-potentiometric characterization of a trans-OM icosa-heme complex, MtrCAB, and demonstrate its capacity to move electrons across a lipid bilayer after incorporation into proteoliposomes. We also show that a stable MtrAB subcomplex can assemble in the absence of MtrC; an MtrBC subcomplex is not assembled in the absence of MtrA; and MtrA is only associated to the membrane in cells when MtrB is present. We propose a model for the modular organization of the MtrCAB complex in which MtrC is an extracellular element that mediates electron transfer to extracellular substrates and MtrB is a trans-OM spanning β-barrel protein that serves as a sheath, within which MtrA and MtrC exchange electrons. We have identified the MtrAB module in a range of bacterial phyla, suggesting that it is widely used in electron exchange with the extracellular environment.

Proceedings of the National Academy of Sciences of the United States of America

95113304

10.1016/J.MSEC.2004.12.002

A branched material based on biomimetic design: Synthesis and electrochemical properties

Based on the biomimetic route the metal material with tree-like fractal structure was prepared, which showed a rough surface observed by scanning electron microscopy. According to the electrochemical experiments, the formed material exhibited the strong catalytic capability as an electrode in hydrogen evolution reaction, comparing with other conventional structure materials under comparable scales. We suggested this promotion of property arose from the contribution of the great surface area and the excellent connectivity offered from the fractal structure. (c) 2005 Elsevier B.V. All rights reserved.

Materials Science and Engineering: C

83938327

10.1007/S002270000466

A functional interpretation of cilia and mucocyte distributions on the abfrontal surface of bivalve gills

Abstract The lack of fundamental data on the abfrontal surface of bivalve gills has prompted a comparative study of cilia and mucocytes on this surface, using scanning electron microscopy and histology on eight species of bivalves, representing seven families and the four major gill types: Mytilus edulis, Modiolus modiolus, Arca zebra, Placopecten magellanicus, Crassostrea virginica, Spisula solidissima, Mercenaria mercenaria, and Mya arenaria. Abfrontal cilia and mucocytes were found in all species studied, with types and densities differing within and between gill types. The three species of homorhabdic filibranchs presented different densities of abfrontal cilia and mucocytes, from very dense in M. edulis to sparse in A. zebra. The heterorhabdic gills had intermediate cilia and mucocyte densities, with highest concentrations of both abfrontal cilia and mucocytes on the principal filaments. The eulamellibranchs showed low ciliary densities together with high mucocyte densities, especially in S. solidissima, where the abfrontal mucocytes were glandular. These results indicate that: (1) the abfrontal surface is a vestigial mucociliary surface; (2) the abfrontal surface cannot participate in water pumping in most species, due to low ciliary densities; and (3) species with high densities of abfrontal mucocytes could utilize abfrontal mucus to reduce drag, especially in the highly fused gills, such as those of the eulamellibranchs. The differing distributions of abfrontal cilia and mucocytes may reflect different selective pressures acting on the gills within the various taxa.

Marine Biology

34132579

10.1126/SCIENCE.1141752

FT Protein Movement Contributes to Long-Distance Signaling in Floral Induction of Arabidopsis

In plants, seasonal changes in day length are perceived in leaves, which initiate long-distance signaling that induces flowering at the shoot apex. The identity of the long-distance signal has yet to be determined. In Arabidopsis, activation of FLOWERING LOCUS T (FT) transcription in leaf vascular tissue (phloem) induces flowering. We found that FT messenger RNA is required only transiently in the leaf. In addition, FT fusion proteins expressed specifically in phloem cells move to the apex and move long distances between grafted plants. Finally, we provide evidence that FT does not activate an intermediate messenger in leaves. We conclude that FT protein acts as a long-distance signal that induces Arabidopsis flowering.

Science

3999677

10.1016/J.SCITOTENV.2017.06.173

Using ecosystem engineers as tools in habitat restoration and rewilding: beaver and wetlands.

Potential for habitat restoration is increasingly used as an argument for reintroducing ecosystem engineers. Beaver have well known effects on hydromorphology through dam construction, but their scope to restore wetland biodiversity in areas degraded by agriculture is largely inferred. Our study presents the first formal monitoring of a planned beaver-assisted restoration, focussing on changes in vegetation over 12years within an agriculturally-degraded fen following beaver release, based on repeated sampling of fixed plots. Effects are compared to ungrazed exclosures which allowed the wider influence of waterlogging to be separated from disturbance through tree felling and herbivory. After 12years of beaver presence mean plant species richness had increased on average by 46% per plot, whilst the cumulative number of species recorded increased on average by 148%. Heterogeneity, measured by dissimilarity of plot composition, increased on average by 71%. Plants associated with high moisture and light conditions increased significantly in coverage, whereas species indicative of high nitrogen decreased. Areas exposed to both grazing and waterlogging generally showed the most pronounced change in composition, with effects of grazing seemingly additive, but secondary, to those of waterlogging. Our study illustrates that a well-known ecosystem engineer, the beaver, can with time transform agricultural land into a comparatively species-rich and heterogeneous wetland environment, thus meeting common restoration objectives. This offers a passive but innovative solution to the problems of wetland habitat loss that complements the value of beavers for water or sediment storage and flow attenuation. The role of larger herbivores has been significantly overlooked in our understanding of freshwater ecosystem function; the use of such species may yet emerge as the missing ingredient in successful restoration.

Science of The Total Environment

88120147

10.2307/1310784

Alteration of North American streams by beaver

organic matter in the channel, create and maintain wetlands, modify nutrient cycling and decomposition dynamics, modify the structure and dynamics of the riparian zone, influence the character of water and materials transported downstream, and ultimately influence plant and animal community composition and diversity (Naiman and Melillo 1984, Naiman et al. 1986). In addition to their importance at the ecosystem level, these effects have a significant impact on the landscape and must be interpreted over broad spatial and temporal scales as beaver population dynamics shift in response to disturbance, food supply, disease, and predation. Although once more prevalent than they are today, beaver-induced alterations to drainage networks are not localized or unusual. Where beaver

BioScience

7372531

10.1007/S00359-010-0522-8

Interaction of liquid epicuticular hydrocarbons and tarsal adhesive secretion in Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae)

Species of various insect orders possess specialised tarsal adhesive structures covered by a thin liquid film, which is deposited in the form of footprints. This adhesive liquid has been suggested to be chemically and physiologically related to the epicuticular lipid layer, which naturally covers the body of insects and acts as the prime barrier to environmental stresses, such as desiccation. The functional efficiency of the layer, however, is jeopardised by partial melting that may occur at physiological temperatures. In this study, light microscopic images of elytral prints show that the epicuticular lipid layer of the Colorado potato beetle Leptinotarsa decemlineata actually is partially liquid and chemical investigations reveal the high similarity of the epicuticular hydrocarbon pattern and the tarsal liquid. By means of chemical manipulation of the surface hydrocarbon composition of live beetles, the substance exchange between their tarsal adhesive hairs and the body surface is monitored. Histological sections of L. decemlineata tarsi, furthermore, reveal glandular cells connected to individual adhesive setae and departing from these results, an idea of a general mechanism of tarsal secretion is developed and discussed in a functional–ecological context.

Journal of Comparative Physiology A-neuroethology Sensory Neural and Behavioral Physiology

12557212

10.1073/PNAS.0812533106

The mechanics of slithering locomotion

In this experimental and theoretical study, we investigate the slithering of snakes on flat surfaces. Previous studies of slithering have rested on the assumption that snakes slither by pushing laterally against rocks and branches. In this study, we develop a theoretical model for slithering locomotion by observing snake motion kinematics and experimentally measuring the friction coefficients of snakeskin. Our predictions of body speed show good agreement with observations, demonstrating that snake propulsion on flat ground, and possibly in general, relies critically on the frictional anisotropy of their scales. We have also highlighted the importance of weight distribution in lateral undulation, previously difficult to visualize and hence assumed uniform. The ability to redistribute weight, clearly of importance when appendages are airborne in limbed locomotion, has a much broader generality, as shown by its role in improving limbless locomotion.

Proceedings of the National Academy of Sciences of the United States of America

183514

10.1242/JEB.038075

Deployment of hagfish slime thread skeins requires the transmission of mixing forces via mucin strands

SUMMARY Hagfishes are benthic marine protovertebrates that secrete copious quantities of slime when threatened. The slime originates as a two-component glandular exudate comprised of coiled bundles of cytoskeletal intermediate filaments (thread skeins) and mucin vesicles. Holocrine secretion of the slime into seawater results in the rapid deployment of both fibrous and mucin components, resulting in about a liter of dilute slime. Deployment of the thread skeins involves their unraveling in a fraction of a second from a 150 μm-long ellipsoid bundle to a thread that is 100× longer. We hypothesized that thread skein deployment requires both vigorous hydrodynamic mixing and the presence of mucin vesicles, both of which are required for whole slime deployment. Here we provide evidence that mixing and mucin vesicles are indeed crucial for skein unraveling. Specifically, we show that mucin vesicles mixed into seawater swell and elongate into high-aspect ratio mucin strands that attach to the thread skeins, transmit hydrodynamic forces to them and effect their unraveling by loading them in tension. Our discovery of mucin strands in hagfish slime not only provides a mechanism for the rapid deployment of thread skeins in vivo, it also helps explain how hagfish slime is able to trap such impressive volumes of seawater via viscous entrainment. We believe that the deployment of thread skeins via their interaction with shear-elongated mucins represents a unique mechanism in biology and may lead to novel technologies for transmitting hydrodynamic forces to microscale particles that would typically be immune to such forces.

The Journal of Experimental Biology

13684160

10.1038/S41598-018-25227-9

Energy and time optimal trajectories in exploratory jumps of the spider Phidippus regius

Jumping spiders are proficient jumpers that use jumps in a variety of behavioural contexts. We use high speed, high resolution video to measure the kinematics of a single regal jumping spider for a total of 15 different tasks based on a horizontal gap of 2–5 body lengths and vertical gap of +/−2 body lengths. For short range jumps, we show that low angled trajectories are used that minimise flight time. For longer jumps, take-off angles are steeper and closer to the optimum for minimum energy cost of transport. Comparison of jump performance against other arthropods shows that Phidippus regius is firmly in the group of animals that use dynamic muscle contraction for actuation as opposed to a stored energy catapult system. We find that the jump power requirements can be met from the estimated mass of leg muscle; hydraulic augmentation may be present but appears not to be energetically essential.

Scientific Reports

87343709

10.2307/1364635

SOCIAL BEHAVIOR IN NESTING CLIFF SWALLOWS

The Condor

6551747

10.1016/S0021-9290(03)00130-1

Understanding stress concentration about a nutrient foramen.

We investigated the microstructural basis of a reduced stress concentration around the primary nutrient foramen of the equine third metacarpus. We quantified the spatial variations of compositional parameters (mineral content, volume fraction, histological architecture, and osteonal trajectories) from microradiographs and polarizing microscopic images of thin sections. These variations in composition and organization in turn cause variations in mechanical properties of cortical bone. We modeled the spatially inhomogeneous anisotropic elastic properties based on the measured compositional parameters and used the properties as inputs to a finite element model of the bone containing the foramen. This model, spatially constructed solely from the microscopic images, was subsequently validated by our mechanical test results. We found that: (1) a primary mechanism for stress concentration reduction appears to be due to an increased compliance near the foramen: the sharp discontinuity represented by the hole is softened by embedding it in a compliant region; (2) a reinforcing ring of increased stiffness exists at some distance from the foramen; and (3) a ring of lamellar bone exists along the foramen inside edge, which might serve to reduce the chance of cracks forming there. Our work is allowing us to design biomimetic structures with holes by mimicking the microstructure near the nutrient foramen.

Journal of Biomechanics

86729322

10.1139/Z99-178

Mechanism, kinematic performance, and fitness consequences of jumping behavior in entomopathogenic nematodes (Steinernema spp.)

The mechanism enabling entomopathogenic nematodes (Steinernema spp.) to jump is described. Jumping performance is measured and the contribution of jumping to host finding is estimated. We used the entomopathogenic nematode Steinernema carpocapsae as a model species for the genus. Nematodes jump using a two-step process of forming and contracting a loop. During loop formation, the nematode bends the anterior half of its body until the head region makes contact with the side of the body. The two body regions are held fast by the surface tension of the water film covering the nematode. When the loop is contracted, the body becomes contorted so that the cuticle kinks. This extreme bending generates and stores sufficient energy that when the surface-tension force is broken the nematode is propelled through the air. The nematode (0.558 mm in length) can jump a distance of 4.8 ± 0.8 mm (mean ± SEM) and a height of 3.9 ± 0.1 mm. The contribution of jumping to host finding varies among species and is related to th...

Canadian Journal of Zoology

4387522

10.1038/NATURE08698

FOXO-dependent regulation of innate immune homeostasis

The innate immune system represents an ancient host defence mechanism that protects against invading microorganisms. An important class of immune effector molecules to fight pathogen infections are antimicrobial peptides (AMPs) that are produced in plants and animals. In Drosophila, the induction of AMPs in response to infection is regulated through the activation of the evolutionarily conserved Toll and immune deficiency (IMD) pathways. Here we show that AMP activation can be achieved independently of these immunoregulatory pathways by the transcription factor FOXO, a key regulator of stress resistance, metabolism and ageing. In non-infected animals, AMP genes are activated in response to nuclear FOXO activity when induced by starvation, using insulin signalling mutants, or by applying small molecule inhibitors. AMP induction is lost in foxo null mutants but enhanced when FOXO is overexpressed. Expression of AMP genes in response to FOXO activity can also be triggered in animals unable to respond to immune challenges due to defects in both the Toll and IMD pathways. Molecular experiments at the Drosomycin promoter indicate that FOXO directly binds to its regulatory region, thereby inducing its transcription. In vivo studies in Drosophila, but also studies in human lung, gut, kidney and skin cells indicate that a FOXO-dependent regulation of AMPs is evolutionarily conserved. Our results indicate a new mechanism of cross-regulation of metabolism and innate immunity by which AMP genes can be activated under normal physiological conditions in response to the oscillating energy status of cells and tissues. This regulation seems to be independent of the pathogen-responsive innate immunity pathways whose activation is often associated with tissue damage and repair. The sparse production of AMPs in epithelial tissues in response to FOXO may help modulating the defence reaction without harming the host tissues, in particular when animals are suffering from energy shortage or stress.

Nature

17762026

10.1242/JEB.00859

The effect of molluscan glue proteins on gel mechanics

SUMMARY Several molluscs have been shown to alternate between a non-adhesive trail mucus and a similar gel that forms a strong glue. The major structural difference between the two secretions is the presence of specific proteins in the adhesive mucus. The present study identifies similar proteins from the glue of the slug Arion subfuscus and the land snail Helix aspersa. To investigate the role played by these proteins in adhesion, the proteins were isolated from the adhesive mucus of different molluscs and added to commercial polymer solutions. The effect was observed qualitatively, and quantified using a dynamic rheometer. The isolated proteins triggered gelling or visible stiffening of agar, pectin and polygalacturonic acid. The effect was stronger on more negatively charged polymers. The effect of the proteins was concentration dependent with an optimal concentration of 1–1.5 mg ml–1, and was weakened when their structure changed. Other proteins and carbohydrates found in the adhesive mucus had no clear mechanical effect on gels. These findings show that the addition of these proteins to large, anionic polymers plays a central role in the formation of a glue from a mucus-like secretion. Such a mechanism may be common among invertebrates, and it may guide biomimetic approaches in the development of glues and gels.

The Journal of Experimental Biology

24405788

10.1016/0040-8166(91)90043-S

Ultrastructure and permeability characteristics of the membranes of mucous granules of the hagfish.

Mucous granules secreted by the slime glands of the hagfish. Eptatretus stouti, were studied after ultrarapid cryofixation and freeze substitution in diverse media (osmium tetroxide in acetone; several aqueous glutaraldehyde-based media with or without osmium). Only freeze substitution with osmium tetroxide-acetone preserved the granules intact, allowing visualization of its single unit membrane. Tests of the rupture or stability of freshly secreted mucous granules in sea water and other aqueous media showed the membranes of the granules are permeable to all inorganic cations tested, ranging in relative mass from ammonium to barium. They are permeable to the univalent anions chloride, nitrate, and bicarbonate, but not to the di- or trivalent anions sulfate, phosphate, and citrate. Moreover, in solutions where nonpenetrant anions were present, rupture occurred if the osmotic pressure was below a critical level (about 800 mOsmol/l). The structural and permeability characteristics of the granules account for the explosive speed with which they rupture, releasing their mucous contents, on contact with sea water.

Tissue & Cell

20009561

10.1093/AOB/MCR307

Restoration of tensile strength in bark samples of Ficus benjamina due to coagulation of latex during fast self-healing of fissures.

BACKGROUND AND AIMS The functions of plant latex have been discussed for a long time. Today, many studies support a defence mechanism as being its main function. A role as a self-healing mechanism was never attributed to the coagulation of latex. In this study we quantified the contribution of the coagulation of Ficus benjamina (weeping fig) latex to a restoration of the mechanical properties of the bark after external lesions. METHODS Tensile tests of F. benjamina bark were conducted either immediately after injury or at various latency times after injury. KEY RESULTS A significant increase in the tensile strength of bark samples until 30 min after injury was found, and this effect could be attributed to the coagulation of plant latex alone. The tensile strength remains nearly constant until several hours or days after injury. Then, very probably due to other mechanisms such as cell growth and cell proliferation, the tensile strength begins to increase slightly again. CONCLUSIONS The coagulation of latex seals lesions and serves as a quick and effective pre-step of subsequent, more effective, long-lasting self-healing mechanisms such as cell growth and proliferation. Thus, a fast self-healing effect can be included in the list of functions of plant latex.

Annals of Botany

53131393

10.1111/J.1095-8312.2009.01362.X

How do sucker-footed bats hold on, and why do they roost head-up?

Individuals of most bat species hang head-down by their toenails from rough surfaces, but Madagascar’s endemic sucker-footed bat (Myzopoda aurita) clings head-up to smooth leaves using specialized pads on its wrists and ankles. We investigated the adhesive performance of 28 individuals and found that attachment performance on brass was not affected by the presence or absence of a seal around the pad–surface interface. Furthermore, on smooth acrylic, the wrist pads were more than nine-fold weaker when lifted perpendicular to the surface than when pulled parallel to it. The unimportance of a seal and the difference in strength in those directions on a smooth surface are characteristic of wet adhesion, but not of suction. Thus, despite its name, the sucker-footed bat appears to adhere using wet adhesion. We observed that when wrist pads were pushed anteriorly, they unpeeled easily from the surface because of deformation of the pads. This most likely permits rapid detachment during crawling, but would also cause passive detachment if bats roosted head-down. This provides an ecomorphological explanation to the head-up roosting behaviour of these unique bats. The results obtained in the present study thus link morphology, behaviour, and roosting ecology for an enigmatic Malagasy endemic. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 233–240. ADDITIONAL KEYWORDS: detachment – ecomorphology – head-up posture – Myzopoda – roosting posture – suction – Thyroptera – wet adhesion.

Biological Journal of The Linnean Society

71716830

10.1371/JOURNAL.PONE.0213071

Companion planting with French marigolds protects tomato plants from glasshouse whiteflies through the emission of airborne limonene

Horticulturalists and gardeners in temperate regions often claim that planting marigolds next to tomato plants protects the tomatoes from the glasshouse whitefly (Trialeurodes vaporariorum Westwood). If shown to hold true, this technique could be used in larger-scale tomato production, protecting the crop and helping to introduce greater plant diversity into these agro-ecosystems. Here we present two large-scale glasshouse trials corresponding to the two main ways growers are likely to use marigolds to control whiteflies. In the first, marigolds are grown next to tomato throughout the growing period and we quantify whitefly population growth from the seedling stage over a 48 day infestation period. Here we show that association with marigolds significantly slows whitefly population development. Introducing additional whitefly-attractive ‘pull’ plants around the perimeter of plots has little effect, but reducing the proportion of marigolds and introducing other non-hosts of whiteflies (basil, nasturtium and Chinese cabbage) also reduces whitefly populations on tomato. The second experiment assesses the efficacy of marigolds when used as an ‘emergency’ measure. Here we allow whitefly populations to build to a high density on unprotected tomatoes then introduce marigolds and assess whitefly population over a further period. Following laboratory work showing limonene to be a major chemical component of French marigolds and a negative behaviour response of whiteflies to this compound, limonene dispensers are added as an additional treatment to this experiment. “Emergency” marigold companion planting yielded minimal reductions in whitefly performance, but the use of limonene dispensers was more effective. Our work indicates that companion planting short vine tomatoes with French marigolds throughout the growing season will slow development of whitefly populations. Introducing marigolds to unprotected tomatoes after significant whitefly build-up will be less effective. The use of limonene dispensers placed near to tomato plants also shows promise. It is argued that this work supports the possibility of the development of a mixture of tomato companion plants that infer ‘associational resistance’ against many major invertebrate pests of tomato. Such a mixture, if comprising edible or ornamental plants, would be economically viable, would reduce the need for additional chemical and biological control, and, if used outdoors, would generate plant-diverse agro-ecosystems that are better able to harbour invertebrate wildlife.

PLOS ONE

21786477

10.1023/A:1022421515027

The Calvin cycle revisited

The sequence of reactions in the Calvin cycle, and the biochemical characteristics of the enzymes involved, have been known for some time. However, the extent to which any individual enzyme controls the rate of carbon fixation has been a long standing question. Over the last 10 years, antisense transgenic plants have been used as tools to address this and have revealed some unexpected findings about the Calvin cycle. It was shown that under a range of environmental conditions, the level of Rubisco protein had little impact on the control of carbon fixation. In addition, three of the four thioredoxin regulated enzymes, FBPase, PRKase and GAPDH, had negligible control of the cycle. Unexpectedly, non-regulated enzymes catalysing reversible reactions, aldolase and transketolase, both exerted significant control over carbon flux. Furthermore, under a range of growth conditions SBPase was shown to have a significant level of control over the Calvin cycle. These data led to the hypothesis that increasing the amounts of these enzymes may lead to an increase in photosynthetic carbon assimilation. Remarkably, photosynthetic capacity and growth were increased in tobacco plants expressing a bifunctional SBPase/FBPase enzyme. Future work is discussed which will further our understanding of this complex and important pathway, particularly in relation to the mechanisms that regulate and co-ordinate enzyme activity.

Photosynthesis Research

7901888

10.1529/BIOPHYSJ.105.069344

Sacrificial bonds and hidden length: unraveling molecular mesostructures in tough materials.

Sacrificial bonds and hidden length in structural molecules and composites have been found to greatly increase the fracture toughness of biomaterials by providing a reversible, molecular-scale energy-dissipation mechanism. This mechanism relies on the energy, of order 100 eV, needed to reduce entropy and increase enthalpy as molecular segments are stretched after being released by the breaking of weak bonds, called sacrificial bonds. This energy is relatively large compared to the energy needed to break the polymer backbone, of order a few eV. In many biological cases, the breaking of sacrificial bonds has been found to be reversible, thereby additionally providing a "self-healing" property to the material. Due to the nanoscopic nature of this mechanism, single molecule force spectroscopy using an atomic force microscope has been a useful tool to investigate this mechanism. Especially when investigating natural molecular constructs, force versus distance curves quickly become very complicated. In this work we propose various types of sacrificial bonds, their combination, and how they appear in single molecule force spectroscopy measurements. We find that by close analysis of the force spectroscopy curves, additional information can be obtained about the molecules and their bonds to the native constructs.

Biophysical Journal

5034723

10.1088/1367-2630/15/12/125022

The origin of traveling waves in an emperor penguin huddle

Emperor penguins breed during the Antarctic winter and have to endure temperatures as low as 50 C and wind speeds of up to 200kmh 1 . To conserve energy, they form densely packed huddles with a triangular lattice structure. Video recordings from previous studies revealed coordinated movements in regular wave-like patterns within these huddles. It is thought that these waves are triggered by individual penguins that locally disturb the huddle structure, and that the traveling wave serves to remove the lattice defects and restore order. The mechanisms that govern wave propagation are currently unknown, however. Moreover, it is unknown if the waves are always triggered by the same penguin in a huddle. Here, we present a model in which the observed wave patterns emerge from simple rules involving only the interactions between directly neighboring individuals, similar to the interaction rules found in other jammed systems, e.g. between cars in a traffic jam. Our model predicts that a 6 Authors to whom any correspondence should be addressed.

New Journal of Physics

452161

10.3389/FMICB.2016.01836

The Interaction between Plants and Bacteria in the Remediation of Petroleum Hydrocarbons: An Environmental Perspective

Widespread pollution of terrestrial ecosystems with petroleum hydrocarbons (PHCs) has generated a need for remediation and, given that many PHCs are biodegradable, bio- and phyto-remediation are often viable approaches for active and passive remediation. This review focuses on phytoremediation with particular interest on the interactions between and use of plant-associated bacteria to restore PHC polluted sites. Plant-associated bacteria include endophytic, phyllospheric, and rhizospheric bacteria, and cooperation between these bacteria and their host plants allows for greater plant survivability and treatment outcomes in contaminated sites. Bacterially driven PHC bioremediation is attributed to the presence of diverse suites of metabolic genes for aliphatic and aromatic hydrocarbons, along with a broader suite of physiological properties including biosurfactant production, biofilm formation, chemotaxis to hydrocarbons, and flexibility in cell-surface hydrophobicity. In soils impacted by PHC contamination, microbial bioremediation generally relies on the addition of high-energy electron acceptors (e.g., oxygen) and fertilization to supply limiting nutrients (e.g., nitrogen, phosphorous, potassium) in the face of excess PHC carbon. As an alternative, the addition of plants can greatly improve bioremediation rates and outcomes as plants provide microbial habitats, improve soil porosity (thereby increasing mass transfer of substrates and electron acceptors), and exchange limiting nutrients with their microbial counterparts. In return, plant-associated microorganisms improve plant growth by reducing soil toxicity through contaminant removal, producing plant growth promoting metabolites, liberating sequestered plant nutrients from soil, fixing nitrogen, and more generally establishing the foundations of soil nutrient cycling. In a practical and applied sense, the collective action of plants and their associated microorganisms is advantageous for remediation of PHC contaminated soil in terms of overall cost and success rates for in situ implementation in a diversity of environments. Mechanistically, there remain biological unknowns that present challenges for applying bio- and phyto-remediation technologies without having a deep prior understanding of individual target sites. In this review, evidence from traditional and modern omics technologies is discussed to provide a framework for plant–microbe interactions during PHC remediation. The potential for integrating multiple molecular and computational techniques to evaluate linkages between microbial communities, plant communities and ecosystem processes is explored with an eye on improving phytoremediation of PHC contaminated sites.

Frontiers in Microbiology

97814460

10.1016/J.SOLMAT.2008.08.009

Optical solar energy adaptations and radiative temperature control of green leaves and tree barks

Trees have adapted to keep leaves and barks cool in sunshine and can serve as interesting bionic model systems for radiative cooling. Silicon solar cells, on the other hand, loose up to one third of their energy efficiency due to heating in intensive sunshine. It is shown that green leaves minimize absorption of useful radiation and allow efficient infrared thermal emission. Since elevated temperatures are detrimental for tensile water flow in the Xylem tissue below barks, the optical properties of barks should also have evolved so as to avoid excessive heating. This was tested by performing optical studies with tree bark samples from representative trees. It was found that tree barks have optimized their reflection of incoming sunlight between 0.7 and 2 μm. This is approximately the optical window in which solar light is transmitted and reflected by green vegetation. Simultaneously, the tree bark is highly absorbing and thus radiation emitting between 6 and 10 μm. These two properties, mainly provided by tannins, create optimal conditions for radiative temperature control. In addition, tannins seem to have adopted a function as mediators for excitation energy towards photo-antioxidative activity for control of radiation damage. The results obtained are used to discuss challenges for future solar cell optimization.

Solar Energy Materials and Solar Cells

12543589

10.1111/J.1556-4029.2011.01773.X

K9 Water Searches: Scent and Scent Transport Considerations

Abstract:  Increased use of water search dogs for detecting submerged bodies has created the need for a better understanding of scent emanating from the bodies and how it transits the water to the dog’s nose. A review of recent literature identifies likely scent sources, potential scent transport processes, and research needs. Scent sources include gases in bubbles or dissolved in the water, liquids as buoyant plumes and droplets or dissolved in the water, and solids consisting of buoyant particulates with secretions, bacteria, and body fluids. Potential transport processes through the water include buoyancy, entrainment, and turbulence. Transport processes from the water surface into the air include volatilization and evaporation enhanced by bubble bursting, breaking waves, splashing, and wind spray. Implications for the use of water search dogs are examined. Observations of submerged, decomposing bodies are needed to quantify the physical and chemical characteristics of the scent and scent transport processes.

Journal of Forensic Sciences

17484445

10.1021/NN200107U

Flaw tolerance of nuclear intermediate filament lamina under extreme mechanical deformation.

The nuclear lamina, composed of intermediate filaments, is a structural protein meshwork at the nuclear membrane that protects genetic material and regulates gene expression. Here we uncover the physical basis of the material design of nuclear lamina that enables it to withstand extreme mechanical deformation of >100% strain despite the presence of structural defects. Through a simple in silico model we demonstrate that this is due to nanoscale mechanisms including protein unfolding, alpha-to-beta transition, and sliding, resulting in a characteristic nonlinear force-extension curve. At the larger microscale this leads to an extreme delocalization of mechanical energy dissipation, preventing catastrophic crack propagation. Yet, when catastrophic failure occurs under extreme loading, individual protein filaments are sacrificed rather than the entire meshwork. This mechanism is theoretically explained by a characteristic change of the tangent stress-strain hardening exponent under increasing strain. Our results elucidate the large extensibility of the nuclear lamina within muscle or skin tissue and potentially many other protein materials that are exposed to extreme mechanical conditions, and provide a new paradigm toward the de novo design of protein materials by engineering the nonlinear stress-strain response to facilitate flaw-tolerant behavior.

ACS Nano

231604

10.1073/PNAS.1106904108

High-speed microscopic imaging of flagella motility and swimming in Giardia lamblia trophozoites

We report, in this paper, several findings about the swimming and attachment mechanisms of Giardia lamblia trophozoites. These data were collected using a combination of a high-contrast CytoViva imaging system and a particle image velocimetry camera, which can capture images at speeds greater than 800 frames/s. Using this system, we discovered that, during rapid swimming of Giardia trophozoites, undulations of the caudal region contributed to forward propulsion combined with the beating of the flagella pairs. It was also discovered, in contrast to previous studies with 10 times slower image sampling technique, that the anterior and posterolateral flagella beat with a clearly defined power stroke and not symmetrical undulations. During the transition from free swimming to attachment, trophozoites modified their swimming behavior from a rapid rotating motion to a more stable planar swimming. While using this planar swimming motion, the trophozoites used the flagella for propulsion and directional control. In addition to examination of the posterolateral and anterior flagella, a model to describe the motion of the ventral flagella was derived, indicating that the ventral flagella beat in an expanding sine wave. In addition, the structure of the ventrocaudal groove creates boundary conditions that determine the form of beating of the ventral flagella. The results from this study indicate that Giardia is able to simultaneously generate both ciliary beating and typical eukaryotic flagellar beating using different pairs of flagella.

Proceedings of the National Academy of Sciences of the United States of America

207169759

10.1242/JEB.055996

Humidity affects the extensibility of an orb-weaving spider's viscous thread droplets

SUMMARY The prey-capture threads found in most spider orb webs rely on viscous droplets for their stickiness. Each droplet is formed of a central mass of viscoelastic glycoprotein glue surrounded by an aqueous covering, both of which incorporate hydrophilic components. We found that the extensibility of droplets on Larinioides cornutus threads increased as humidity increased. However, the deflection of the droplets' supporting axial lines did not change, indicating that atmospheric water uptake increases glycoprotein plasticity, but not glycoprotein adhesion. The extensibility of droplets, along with that of the thread's supporting axial fibers, is responsible for summing the adhesion of multiple thread droplets. Therefore, daily changes in humidity have the potential to significantly alter the performance of viscous threads and orb webs.

The Journal of Experimental Biology

6587094

10.1086/380213

Lipids of the Stratum Corneum Vary with Cutaneous Water Loss among Larks along a Temperature‐Moisture Gradient

We explored the relationship between lipids of the stratum corneum (SC), the barrier to water‐vapor diffusion of the skin, and cutaneous water loss (CWL) of species of free‐living larks along a temperature‐moisture gradient. Our results showed that free fatty acids, cholesterol, and ceramides were the major constituents of SC in larks from different environments including the Netherlands, a mesic environment; Iran, a semiarid region; and several areas in Saudi Arabia, a hot dry desert. We found that CWL was reduced among larks inhabiting deserts, but our data did not support the hypothesis that birds from desert environments have larger quantities of lipids per unit dry mass of the SC than larks from more mesic environments. Instead, larks in arid environments had a higher proportion of ceramides, especially the more polar fractions 4–6, and a smaller proportion of free fatty acids in their SC, an adjustment that apparently reduced their CWL. Subtle changes in the ratios of lipid classes can apparently alter the movement of water vapor through the skin. We hypothesize that desert birds have higher proportions of ceramides in their SC and lower proportions of free fatty acids because this combination allows the lipid lamellae to exist in a more highly ordered crystalline phase and consequently creates a tighter barrier to water‐vapor diffusion.

Physiological and Biochemical Zoology

18963609

10.1073/PNAS.0912254107

A dinoflagellate exploits toxins to immobilize prey prior to ingestion

Toxins produced by the harmful algal bloom (HAB) forming, mixotrophic dinoflagellate Karlodinium veneficum have long been associated with fish kills. To date, the perceived ecological role for toxins has been relief from grazing pressures. Here, we demonstrate that karlotoxins also serve as a predation instrument. Using high-speed holographic microscopy, we measure the swimming behavior of several toxic and nontoxic strains of K. veneficum and their prey, Storeatula major, within dense suspensions. The selected strains produce toxins with varying potency and dosages, including a nontoxic one. Results clearly show that mixing the prey with the predatory, toxic strains causes prey immobilization at rates that are consistent with the karlotoxins’ potency and dosage. Even prey cells that continue swimming slow down after exposure to toxic predators. The swimming characteristics of predators vary substantially in pure suspensions, as quantified by their velocity, radii of helical trajectories, and direction of helical rotation. When mixed with prey, all toxic strains that are involved in predation slow down. Furthermore, they substantially reduced their predominantly vertical migration, presumably to remain in the vicinity of their prey. Conversely, the nontoxic control strain does not alter its swimming and does not affect prey behavior. In separate experiments, we show that exposing prey to exogenous toxins also causes prey immobilization at rates consistent with potency. Clearly, the toxic predatory strains use karlotoxins as a means of stunning their prey, before ingesting it. These findings add a substantiated critical understanding for why some HAB species produce such complex toxin molecules.

Proceedings of the National Academy of Sciences of the United States of America

2223462

10.1021/BM049899T

Giant bent-core mesogens in the thread forming process of marine mussels.

In marine mussels (Mytilus), byssal threads are made in minutes from prefabricated smectic polymer liquid crystals by a process resembling reaction injection molding. The mesogens in these arrays are known to be natural block copolymers with rodlike collagen cores. Using atomic force microscopy, it was shown that these collagenous mesogens are bent-core or banana-shaped in a manner that is consistent with and predictable from their amino acid sequence. The overall bend angle in preCOL-NG in Mytilus galloprovincialis is about 130 degrees. The mesogens have a center-to-center separation of approximately 22 nm and a length of 200 nm. It is evident that the smectic structure of the prefabricated mesophases remains largely intact over 1-3 microm distances in the molded fibers and is presumably locked in place during molding by cross-linking. Like the smectic liquid crystals of many synthetic banana mesogens, the collagenous mesogens of the byssal threads exhibit SmC(2) symmetry with a characteristic tilt of 24.6 degrees. At about 100% extension, this tilt is considerably reduced and the globular end domains are no longer visible presumably because they have been unraveled.

Biomacromolecules

46523397

10.1152/AJPLEGACY.1969.217.5.1504

Physiological responses to heat stress in the poorwill.

American Journal of Physiology

87369205

10.2307/1538612

SECRETION OF GASES AGAINST HIGH PRESSURES IN THE SWIMBLADDER OF DEEP SEA FISHES II. THE RETE MIRABILE

1. The structure and dimensions are given for the rete mirabile. It is interpreted as a counter current diffusion exchange mechanism between the afferent and efferent rete.2. This diffusion exchange has been quantitatively evaluated for O2, CO2, and N2. The arrangement makes possible the maintenance of a steep tension gradient within the rete, so that the oxygen loss from the leaving blood can be extremely low.3. Quantitative evaluation of the efficiency of the rete diffusion makes it clear that the limiting equilibrium pressure in the swimbladder could be extremely high, and that the limiting factor lies mainly in the dissociation pressure of the chemical or physical reaction which ultimately splits off the oxygen.4. The nature of the responsible compound and reaction is unknown. The anatomical arrangement of the glandular structure and the rete is suggestive of a cellular secretion of oxygen rather than of a mechanism that splits off oxygen in the blood.5. Data on the deposition of nitrogen against high...

The Biological Bulletin

2469071

10.4161/PSB.21661

Tuning of color contrast signals to visual sensitivity maxima of tree shrews by three Bornean highland Nepenthes species

Three species of Nepenthes pitcher plants (Nepenthes rajah, Nepenthes lowii and Nepenthes macrophylla) specialize in harvesting nutrients from tree shrew excreta in their pitchers. In all three species, nectaries on the underside of the pitcher lid are the focus of the tree shrews' attention. Tree shrews are dichromats, with visual sensitivity in the blue and green wavebands. All three Nepenthes species were shown to produce visual signals, in which the underside of the pitcher lid (the area of highest nectar production) stood out in high contrast to the adjacent area on the pitcher (i.e., was brighter), in the blue and green wavebands visible to the tree shrews. N. rajah showed the tightest degree of “tuning,” notably in the green waveband. Conversely, pitchers of Nepenthes burbidgeae, a typical insectivorous species sympatric with N. rajah, did not produce a color pattern tuned to tree shrew sensitivity maxima.

Plant Signaling & Behavior

23199158

10.1098/RSPB.2004.2850

Adhesion forces measured at the level of a terminal plate of the fly's seta

The attachment pads of fly legs are covered with setae, each ending in small terminal plates coated with secretory fluid. A cluster of these terminal plates contacting a substrate surface generates strong attractive forces that hold the insect on smooth surfaces. Previous research assumed that cohesive forces and molecular adhesion were involved in the fly attachment mechanism. The main elements that contribute to the overall attachment force, however, remained unknown. Multiple local force–volume measurements were performed on individual terminal plates by using atomic force microscopy. It was shown that the geometry of a single terminal plate had a higher border and considerably lower centre. Local adhesion was approximately twice as strong in the centre of the plate as on its border. Adhesion of fly footprints on a glass surface, recorded within 20 min after preparation, was similar to adhesion in the centre of a single attachment pad. Adhesion strongly decreased with decreasing volume of footprint fluid, indicating that the layer of pad secretion covering the terminal plates is crucial for the generation of a strong attractive force. Our data provide the first direct evidence that, in addition to Van der Waals and Coulomb forces, attractive capillary forces, mediated by pad secretion, are a critical factor in the fly's attachment mechanism.

Proceedings of The Royal Society B: Biological Sciences

32170919

10.1007/S005720100108

Exploration types of ectomycorrhizae

Abstract  There is a need to quantify and qualify the ability of ectomycorrhizae to improve tree growth and nutrition and, in particular, to define criteria to classify ectomycorrhizae with respect to ecologically relevant features. Whereas the numbers of ectomycorrhizae and morphotypes give useful information when related to root biomass, root length or soil volume, the development and differentiation of the extramatrical mycelium may represent important predicative features relevant to the ecological classification of ectomycorrhizae. Here, different exploration types of ectomycorrhizae are distinguished based on the amount of emanating hyphae or the presence and differentiation of rhizomorphs. Their putative ecological importance is discussed.

Mycorrhiza

3246541

10.1073/PNAS.1210105109

Pointillist structural color in Pollia fruit

Biological communication by means of structural color has existed for at least 500 million years. Structural color is commonly observed in the animal kingdom, but has been little studied in plants. We present a striking example of multilayer-based strong iridescent coloration in plants, in the fruit of Pollia condensata. The color is caused by Bragg reflection of helicoidally stacked cellulose microfibrils that form multilayers in the cell walls of the epicarp. We demonstrate that animals and plants have convergently evolved multilayer-based photonic structures to generate colors using entirely distinct materials. The bright blue coloration of this fruit is more intense than that of any previously described biological material. Uniquely in nature, the reflected color differs from cell to cell, as the layer thicknesses in the multilayer stack vary, giving the fruit a striking pixelated or pointillist appearance. Because the multilayers form with both helicoidicities, optical characterization reveals that the reflected light from every epidermal cell is polarized circularly either to the left or to the right, a feature that has never previously been observed in a single tissue.

Proceedings of the National Academy of Sciences of the United States of America

3915031

10.1098/RSPB.2016.2569

The acacia ants revisited: convergent evolution and biogeographic context in an iconic ant/plant mutualism

Phylogenetic and biogeographic analyses can enhance our understanding of multispecies interactions by placing the origin and evolution of such interactions in a temporal and geographical context. We use a phylogenomic approach—ultraconserved element sequence capture—to investigate the evolutionary history of an iconic multispecies mutualism: Neotropical acacia ants (Pseudomyrmex ferrugineus group) and their associated Vachellia hostplants. In this system, the ants receive shelter and food from the host plant, and they aggressively defend the plant against herbivores and competing plants. We confirm the existence of two separate lineages of obligate acacia ants that convergently occupied Vachellia and evolved plant-protecting behaviour, from timid ancestors inhabiting dead twigs in rainforest. The more diverse of the two clades is inferred to have arisen in the Late Miocene in northern Mesoamerica, and subsequently expanded its range throughout much of Central America. The other lineage is estimated to have originated in southern Mesoamerica about 3 Myr later, apparently piggy-backing on the pre-existing mutualism. Initiation of the Pseudomyrmex/Vachellia interaction involved a shift in the ants from closed to open habitats, into an environment with more intense plant herbivory. Comparative studies of the two lineages of mutualists should provide insight into the essential features binding this mutualism.

Proceedings of The Royal Society B: Biological Sciences

25556363

10.1007/BF00018716

The behavior in flow of the morphologically variable seaweed Hedophyllum sessile (C. Ag.) Setchell

Drag force was measured on individual specimens of Hedophyllum sessile in a variable-speed flow tank. Those from sheltered localities, which are broad, bullate blades, experience greater drag at a given water velocity than ones from localities more exposed to the action of waves, which have smooth, deeply dissected blades. All specimens rearranged their blades as water velocity increased, resulting in a decrease in effective drag at higher water speeds, but individuals with smooth, dissected blades assumed a more compact shape at high current speeds and thus reduced their effective drag over that of broad-bladed individuals at the same speed. In habitats chronically exposed to strong wave action, drag reduction may be an important survival mechanism; in calm habitats the turbulence induced by lack of such streamlining may enhance mixing of the water in the immediate vicinity of a plant.

Hydrobiologia

84148018

10.1006/JTBI.1995.0219

THE MECHANICS OF SEED EXPULSION IN ACANTHACEAE

Abstract The bilocular seed capsules of species on the Acanthaceae subfamily Acanthoideae are either hygrochastic or xerochastic, but in both cases the mechanism for seed expulsion is similar; only the “trigger” differs in the two instances. The drying of the capsule results in the storage of elastic energy in the capsule valves. The failure of the seam joining the two values precipitates the conversion of the elastic potential energy stored in the valves and seeds. In the hygrochastic case the failure is due to moisture absorption on wetting of the capsule beak which weakens the pectic “glue”; in the xerochastic case the seam failure is due simply to the high stress in the bonding layer at some degree of desiccation. This paper explains quantitatively how the anatomy of the capsule efficiently imports high initial expulsion velocity to the seeds in order to maximize their range. The specific example considered inRuellia brittonianaLeonard, a cultivated shrub native to Mexico, but the situation is similar for the entire Acanthoideae subfamily.

Journal of Theoretical Biology

33913226

10.1093/ICB/ICQ033

Thermal windows on Brazilian free-tailed bats facilitate thermoregulation during prolonged flight.

The Brazilian free-tailed bat (Tadarida brasiliensis) experiences challenging thermal conditions while roosting in hot caves, flying during warm daylight conditions, and foraging at cool high altitudes. Using thermal infrared cameras, we identified hot spots along the flanks of free-ranging Brazilian free-tailed bats, ventral to the extended wings. These hot spots are absent in syntopic cave myotis (Myotis velifer), a species that forages over relatively short distances, and does not engage in long-distance migration. We hypothesized that the hot spots, or "radiators," on Brazilian free-tailed bats may be adaptations for migration, particularly in this long-distance, high-flying species. We examined the vasculature of radiators on Brazilian free-tailed bats with transillumination to characterize the unique arrangements of arteries and veins that are positioned perpendicular to the body in the proximal region of the wing. We hypothesized that these radiators aid in maintaining heat balance by flushing the uninsulated thermal window with warm blood, thereby dissipating heat while bats are flying under warm conditions, but shunting blood away and conserving heat when they are flying in cooler air at high altitudes. We also examined fluid-preserved specimens representing 122 species from 15 of 18 chiropteran families and radiators appeared present only in species in the family Molossidae, including both sedentary and migratory species and subspecies. Thus, the radiator appears to be a unique trait that may facilitate energy balance and water balance during sustained dispersal, foraging, and long-distance migration.

Integrative and Comparative Biology

100969465

10.1007/S10533-005-4279-Z

Nitrogen Mineralization, Ammonia Accumulation, and Emission of Gaseous NH3 by Soil-feeding Termites

There are numerous reports on the accumulation of ammonia in the mounds of soil-feeding termites. Here, we provided direct evidence for an effective mineralization of nitrogenous soil organic matter in the gut of Cubitermes spp., which gives rise to enormous ammonia concentrations in the intestinal tract. In Cubitermes ugandensis, the ammonia content of the nest material [24.5 μmol (g dry wt.)−1] was about 300-fold higher than that of the parent soil. Large amounts of ammonia were present throughout the intestinal tract, with lowest values in the extremely alkaline gut sections (pH >12) and highest values posterior hindgut [185 μmol (g dry wt.)−1]. Results obtained with other Cubitermes species were similar. Ammonia concentrations in the posterior hindgut of these humivorous species (up to 130 mM) are among the highest values ever reported for soil macroinvertebrates and are matched only by insects feeding on an extremely protein-rich diet (e.g., the sarcophageous larvae of blowflies). Volatilization of ammonia [about 10 nmol (g fresh wt.)−1 h−1], either directly by emission from the termite body or indirectly from their feces, led to NH3 concentrations in the nest atmosphere of C. ugandensis that were three orders of magnitude above the ambient background – a relative accumulation that is considerably higher than that observed with CH4 and CO2. Together with previous results, these observations document that through their feeding activity and due to the physicochemical and biochemical properties of their digestive system, soil-feeding termites effectively catalyze the transformation of refractory soil organic nitrogen to a plant-available form that is protected from leaching by adsorption to the nest soil. Nitrogen mineralization rates of soil-feeding termites may surpass those effected by tropical earthworms and should contribute significantly to nitrogen fluxes in tropical ecosystems.

Biogeochemistry

2375135

10.1126/SCIENCE.1174269

Bioluminescence in the Ocean: Origins of Biological, Chemical, and Ecological Diversity

In Living Color Light produced using the chemical process of bioluminescence spans the entire range of the visible spectrum. Bioluminescence has evolved independently several times in the tree of life. However, the majority of bioluminescent organisms reside in the open ocean, where their bioluminescence helps species in over 700 genera evade predators, attract mates, and find food. Widder (p. 704) reviews recent advances in understanding the evolution and distribution of bioluminescence in marine systems. From bacteria to fish, a remarkable variety of marine life depends on bioluminescence (the chemical generation of light) for finding food, attracting mates, and evading predators. Disparate biochemical systems and diverse phylogenetic distribution patterns of light-emitting organisms highlight the ecological benefits of bioluminescence, with biochemical and genetic analyses providing new insights into the mechanisms of its evolution. The origins and functions of some bioluminescent systems, however, remain obscure. Here, I review recent advances in understanding bioluminescence in the ocean and highlight future research efforts that will unite molecular details with ecological and evolutionary relationships.

Science

4425287

10.1038/NATURE03661

Extracellular electron transfer via microbial nanowires

Microbes that can transfer electrons to extracellular electron acceptors, such as Fe(iii) oxides, are important in organic matter degradation and nutrient cycling in soils and sediments. Previous investigations on electron transfer to Fe(iii) have focused on the role of outer-membrane c-type cytochromes. However, some Fe(iii) reducers lack c-cytochromes. Geobacter species, which are the predominant Fe(iii) reducers in many environments, must directly contact Fe(iii) oxides to reduce them, and produce monolateral pili that were proposed, on the basis of the role of pili in other organisms, to aid in establishing contact with the Fe(iii) oxides. Here we report that a pilus-deficient mutant of Geobacter sulfurreducens could not reduce Fe(iii) oxides but could attach to them. Conducting-probe atomic force microscopy revealed that the pili were highly conductive. These results indicate that the pili of G. sulfurreducens might serve as biological nanowires, transferring electrons from the cell surface to the surface of Fe(iii) oxides. Electron transfer through pili indicates possibilities for other unique cell-surface and cell–cell interactions, and for bioengineering of novel conductive materials.

Nature

1882899

10.1242/JEB.00596

Phenotypic flexibility in cutaneous water loss and lipids of the stratum corneum

SUMMARY When vertebrates invaded land during the Carboniferous period, they were exposed not only to new ecological opportunities but also to a desiccating environment. To maintain cellular water homeostasis, natural selection modified the integument of pioneering terrestrial animals, enabling them to reduce water loss through the skin. In desert environments, where ambient temperatures (Ta) can reach 50°C, relative humidities are low and drinking water is scarce, integumentary modifications that reduce cutaneous water loss (CWL) could be fundamental to survival. Previous research has shown that hoopoe larks (Alaemon alaudipes) from the Arabian desert reduced CWL when acclimated to 35°C compared with individuals at 15°C, but skylarks (Alauda arvensis) and woodlarks (Lullula arborea), from The Netherlands, and Dunn's larks (Eremalauda dunni), also from the Arabian desert, did not. Here, we test the idea that hoopoe larks acclimated to 35°C would alter the lipid composition of their stratum corneum (SC), resulting in a decrease in CWL, but that skylarks, woodlarks and Dunn's larks would not. Specifically, we hypothesized that hoopoe larks, acclimated to 35°C, would increase the proportions of polar ceramide content and decrease the proportions of free fatty acids in their SC compared with individuals acclimated to 15°C. Results showed that hoopoe larks at 35°C had lower CWL and higher proportions of total ceramides but lower proportions of free fatty acids and sterols in their SC. We demonstrate that adjustments in ratios of lipid classes in the SC are associated with changes in CWL in hoopoe larks.

The Journal of Experimental Biology

10346844

10.2166/WST.2006.349

Microbial conversion of selected azo dyes and their breakdown products.

Four selected azo dyes (acid orange 6, acid orange 7, methyl orange and methyl red) were completely decolourised in the presence of anaerobic granular sludge, while only methyl red was degraded in aerobic conditions using a conventional activated sludge. Additional experiments with culture broth devoid of cells showed that anaerobic decolourisation of azo dyes was performed by extracellular reducing agents produced by anaerobic bacteria. This was further confirmed by abiotic experiments with sulphide and NADH. The presence of redox mediators such as riboflavin led to dramatic acceleration of the anaerobic biodecolourisation process. The azo dye reduction products were found to be sulphanilic acid and 4-aminoresorcinol for acid orange 6; sulphanilic acid and 1-amino-2-naphthol for acid orange 7; N,N-dimethyl-1,4-phenylenediamine and sulphanilic acid for methyl orange; and N,N-dimethyl-1,4-phenylenediamine and anthranilic acid for methyl red. Anaerobic toxicity assays showed that the azo dyes were more toxic than their breakdown products (aromatic amines), except 1-amino-2-naphthol. In the presence of activated sludge, only anthranilic acid was completely mineralised while sulphanilic acid was persistent. 4-aminoresorcinol, 1-amino-2-naphthol and N,N-dimethyl-1,4-phenylenediamine underwent autooxidation in aerobic conditions yielding coloured polymeric products. On the contrary, in the presence of granular methanogenic sludge, 4-aminoresorcinol, 1-amino-2-naphthol and anthranilic acid were quantitatively methanised, sulphanilic acid was partially (70%) mineralised while N,N-dimethyl-1,4-phenylenediamine was only demethylated producing 1,4-phenylenediamine as an end product.

Water Science and Technology

86340418

10.1111/J.1365-2435.2011.01851.X

Sun and shade leaves of Olea europaea respond differently to plant size, light availability and genetic variation

1. Canopy plasticity, the expression of different leaf phenotypes within the crown of an individual tree has complex functional and evolutionary implications that remain to be thoroughly assessed. We hypothesized that it can lead to disparity in how leaves in different positions of the canopy change with allometric growth and population genetic structure. 2. Leaf phenotypes of the inner and outer canopy were estimated using eight morphological and physiological characters. All traits were measured under field conditions in six populations of Olea europaea and again in a common garden for a subset of the genotypes. The same populations were characterized genetically with amplified fragment length polymorphisms (AFLP) genomic scans. With these data, we investigated the extent to which leaf phenotypes change with plant size, genetic processes and in response to environmental conditions inside and outside the canopy. 3. The size of trees measured in the field was clearly associated with the phenotype of sun but not to that of shade leaves. The phenotype of sun leaves depended on both direct and diffuse light, while that of shade leaves was found to correlate only with diffuse radiation. Additionally, light availability inside the canopy was conditioned by the shape of external leaves, and increasing elongation of sun leaves led to higher radiation in the inner canopy. 4. The field phenotypes of both inner and outer canopy leaves were correlated with genetic variation among populations. Conversely, in the common garden, the different genotypes expressed a homogeneous sun phenotype, while phenotypic differences among populations remained apparent in shade leaves. 5. We conclude that, in agreement with our working hypothesis, canopy plasticity is both cause and consequence of the environment experienced by the plant and might lead to the differential expression of genetic polymorphisms among leaves. Furthermore, we propose that it can contribute to buffer abiotic stress and to the partition of light use within the tree crown.

Functional Ecology

18631592

10.1073/PNAS.1112288108

Knickkopf protein protects and organizes chitin in the newly synthesized insect exoskeleton

During each molting cycle of insect development, synthesis of new cuticle occurs concurrently with the partial degradation of the overlying old exoskeleton. Protection of the newly synthesized cuticle from molting fluid enzymes has long been attributed to the presence of an impermeable envelope layer that was thought to serve as a physical barrier, preventing molting fluid enzymes from accessing the new cuticle and thereby ensuring selective degradation of only the old one. In this study, using the red flour beetle, Tribolium castaneum, as a model insect species, we show that an entirely different and unexpected mechanism accounts for the selective action of chitinases and possibly other molting enzymes. The molting fluid enzyme chitinase, which degrades the matrix polysaccharide chitin, is not excluded from the newly synthesized cuticle as previously assumed. Instead, the new cuticle is protected from chitinase action by the T. castaneum Knickkopf (TcKnk) protein. TcKnk colocalizes with chitin in the new cuticle and organizes it into laminae. Down-regulation of TcKnk results in chitinase-dependent loss of chitin, severe molting defects, and lethality at all developmental stages. The conservation of Knickkopf across insect, crustacean, and nematode taxa suggests that its critical roles in the laminar ordering and protection of exoskeletal chitin may be common to all chitinous invertebrates.

Proceedings of the National Academy of Sciences of the United States of America

20628446

10.1242/JEB.056275

Circumferential spicule growth by pericellular silica deposition in the hexactinellid sponge Monorhaphis chuni

SUMMARY The giant basal spicule of the hexactinellid sponge Monorhaphis chuni represents the longest natural siliceous structure on Earth. This spicule is composed of concentrically arranged lamellae that are approximately 10 μm thick. In the present study, we investigated the formation of outer lamellae on a cellular level using microscopic and spectroscopic techniques. It is shown that the formation of an outermost lamella begins with the association of cell clusters with the surface of the thickening and/or growing spicule. The cells release silica for controlled formation of a lamella. The pericellular (silica) material fuses to a delimited and textured layer of silica with depressions approximately 20–30 μm in diameter. The newly formed layer initially displays 40 μm wide, well-structured banded ribbons and only attains its plain surface in a final step. The chemical composition in the depressions was studied using energy dispersive X-ray spectroscopy and by staining with Texas Red. The data suggest that those depressions are the nests for the silica-forming cells and that silica formation starts with a direct association of silica-forming cells with the outer surface of the spicule, where they remain and initiate the development of the next lamellae.

The Journal of Experimental Biology

24346201

10.1104/PP.108.122606

Regulation of CONSTANS and FLOWERING LOCUS T Expression in Response to Changing Light Quality1[C][OA]

In addition to pathways that regulate flowering in response to environmental signals such as photoperiod or cold temperatures (vernalization), flowering time is also regulated by light quality. In many species, far-red (FR) light is known to accelerate flowering. This is environmentally significant because leaves absorb more red light than FR light; thus, plants growing under a canopy experience light that is enriched in FR light. In this article, we have explored the promotion of flowering by FR-enriched light (FREL) in Arabidopsis (Arabidopsis thaliana). Previous work has shown that the floral promoter CONSTANS (CO) plays a critical role in day-length perception and exhibits complex regulation; CO mRNA is regulated by the circadian clock and CO protein is stabilized by light and degraded in darkness. We find that plants grown under FREL contain higher levels of CO mRNA in the early part of the day than plants under white light. Furthermore, transgenic plants expressing CO under the control of a constitutive promoter accumulate higher levels of CO protein under FREL, indicating that FREL can increase CO protein levels independently of transcription. Consistent with the model that FREL promotes flowering through CO, mutants for co or gigantea, which are required for CO transcript accumulation, are relatively insensitive to FREL. Because the red:FR ratios used in these experiments are in the range of what plants would experience under a canopy, these results indicate that the regulation of CO by light quality likely plays a key role in the regulation of flowering time in natural environments.

Plant Physiology

1617982

10.1371/JOURNAL.PONE.0047867

How Archer Fish Achieve a Powerful Impact: Hydrodynamic Instability of a Pulsed Jet in Toxotes jaculatrix

Archer fish knock down insects anchored to vegetation by hitting them with a precisely aimed jet of water. The striking force of the jet at the impact is such to overcome the strong anchoring forces of insects. The origin of the effectiveness of such hunting mechanism has been long searched for inside of the fish, in the unsuccessful attempt to identify internal structures dedicated to the amplification of muscular power. Here we perform a kinematic analysis of the jet emitted by two specimens of Toxotes jaculatrix. We estimate that at the impact the jet conveys a typical specific power of about 3000 W/kg, which is well above the maximum specific power of the order of 500 W/kg deliverable by a vertebrate muscle. Unexpectedly, we find that the amplification of muscular power occurs outside of the fish, and is due to a hydrodynamic instability of the jet akin to those occurring in Drop-on-Demand inkjet printing. The investigated fish are found to modulate the velocity of the jet at the orifice to favor the formation of a single, large, water drop that hits the prey abruptly with a large momentum. The observed mechanism represents a remarkable example of use of an external hydrodynamic lever that does possibly not entail the high evolutionary cost needed for the development of highly specialized internal structures dedicated to the storing of mechanical energy.

PLOS ONE

81015577

10.1016/S0065-2806(10)38003-9

Diverse Strategies of Protein Sclerotization in Marine Invertebrates: Structure–Property Relationships in Natural Biomaterials

Abstract The conventional wisdom regarding sclerotization in arthropod exoskeletons is that tissue stiffness and polyphenolic content are tightly coupled. Recent progress on the biochemistry and mechanics of sclerotized structures from non-arthropod invertebrates such as mussels, polychaetes and squid suggests that this premise is too simple and needs to be more closely scrutinized. Emerging mechanistic insights about structure–property relationships in sclerotized extracellular composites include the following: (1) tissue stiffness can be significantly adjusted by the relative organization and density of rigid versus flexible domains in the protein components (mussel byssal thread); (2) structures can be rich in polyphenols without being sclerotized (mussel adhesive plaque); (3) stiffness and hardness can be adjusted by the relative abundance of interactions between protein-based imidazole and catecholate ligands and transition metals (polychaete jaws and mussel cuticle); (4) polyphenol-derived covalent cross-link density in a protein–chitin composite offers some stiffening, but gains from incrementally controlled dehydration are far greater (squid beak); and finally, (5) protein domains, ligand and metal densities, covalent cross-linking and dehydration are elaborately manipulated to create stiffness gradients between mechanically mismatched tissues.

Advances in Insect Physiology

32478760

10.1007/BF00317628

“Magnetic” termite mound surfaces are oriented to suit wind and shade conditions

SummaryThe termites Amitermes meridionalis and A. laurensis construct remarkable meridional or “magnetic” mounds in northern Australia. These mounds vary geographically in mean orientation in a manner that suggests such variation is an adaptive response to local environmental conditions. Theoretical modelling of solar irradiance and mound rotation experiments show that maintenance of an eastern face temperature plateau during the dry season is the most likely physical basis for the mound orientation response. Subsequent heat transfer analysis shows that habitat wind speed and shading conditions also affect face temperature gradients such as the rate of eastern face temperature change. It is then demonstrated that the geographic variation in mean mound orientation follows the geographic variation in long-term wind speed and shading conditions across northern Australia such that an eastern face temperature plateau is maintained in all locations.

Oecologia

22410850

10.1126/SCIENCE.1067778

Control of the Selectivity of the Aquaporin Water Channel Family by Global Orientational Tuning

Aquaporins are transmembrane channels found in cell membranes of all life forms. We examine their apparently paradoxical property, facilitation of efficient permeation of water while excluding protons, which is of critical importance to preserving the electrochemical potential across the cell membrane. We have determined the structure of the Escherichia coliaquaglyceroporin GlpF with bound water, in native (2.7 angstroms) and in W48F/F200T mutant (2.1 angstroms) forms, and carried out 12-nanosecond molecular dynamics simulations that define the spatial and temporal probability distribution and orientation of a single file of seven to nine water molecules inside the channel. Two conserved asparagines force a central water molecule to serve strictly as a hydrogen bond donor to its neighboring water molecules. Assisted by the electrostatic potential generated by two half-membrane spanning loops, this dictates opposite orientations of water molecules in the two halves of the channel, and thus prevents the formation of a “proton wire,” while permitting rapid water diffusion. Both simulations and observations revealed a more regular distribution of channel water and an increased water permeability for the W48F/F200T mutant.

Science

37408603

10.1002/JEZ.A.116

Shell growth and chamber formation of aquarium-reared Nautilus pompilius (Mollusca, Cephalopoda) by X-ray analysis.

Observations on the growth rate of aquarium maintained Nautilus pompilius in different developmental stages, i.e. juveniles (shell length about 8.75 cm), late juveniles (approximately 10 cm), and early adolescent (approximately 13.5 cm), indicate that this species is fully grown at an age of 7.3-8 years. The age calculations are based on two different computations: (1) the measurement of the increase of the shell length per day and (2) the formation of new septa in time intervals of 150+/-5 days, as demonstrated by X-ray analyses. After N. pompilius hatches, its shell grows about 139 mm to reach full growth and approximately 28 septa are formed. With an increase of the shell length of 0.052 mm per day, it takes about 2,673 days (7.3 years) to reach maturity. Provided that the process of chamber formation follows an exponential function, these computations result in approximately 2,925 days (8 years) to reach full maturity. Supposing that N. pompilius may live for several years after onset of maturity like Nautilus belauensis, the total life span for this species may exceed 11-12 years.

Journal of Experimental Zoology Part A: Comparative Experimental Biology

27129622

10.3732/AJB.0900285

Dehydration-induced expression of a 31-kDa dehydrin in Polypodium polypodioides (Polypodiaceae) may enable large, reversible deformation of cell walls.

Current and predicted climate changes caused by global warming compel greater understanding of the molecular mechanisms that plants use to survive drought. The desiccation-tolerant fern Polypodium polypodioides exhibits extensive cell wall folding when dried to less than 15% relative water content (RWC) and rapidly (within 24 h) rehydrates when exposed to water and high humidity. A 31-kDa putative dehydrin polypeptide expressed in partially and fully dry tissues detected via western blotting was present only during drying and rapidly dissipated (within 24 h) upon tissue rehydration. Immunostaining indicates the presence of dehydrin near the cell wall of partially and fully dried tissues. Atomic force microscopy of tracheal scalariform perforations indicates that dry vascular tissue does not undergo significant strain. Additionally, environmental scanning electron microscopy reveals differential hydrophilicity between the abaxial and adaxial leaf surfaces as well as large, reversible deformation. The ability to avoid cell wall damage in some desiccation-tolerant species may be partially attributed to cell wall localization of dehydrins enabling reversible, large cell-wall deformation. Thus, the de novo synthesis of dehydrin proteins and potential localization to the cell walls of these desiccation-tolerant species may play a role in avoiding mechanical failure during drought.

American Journal of Botany

25939121

10.1080/00218460902782188

Halogenated DOPA in a Marine Adhesive Protein

The sandcastle worm Phragmatopoma californica, a marine polychaete, constructs a tube-like shelter by cementing together sand grains using a glue secreted from the building organ in its thorax. The glue is a mixture of post-translationally modified proteins, notably the cement proteins Pc-1 and Pc-2 with the amino acid, 3,4-dihydroxyphenyl-L-alanine (DOPA). Significant amounts of a halogenated derivative of DOPA were isolated from the worm cement following partial acid hydrolysis and capture of catecholic amino acids by phenylboronate affinity chromatography. Analysis by tandem mass spectrometry and 1H NMR indicates the DOPA derivative to be 2-chloro-4, 5-dihydroxyphenyl-L-alanine. The potential roles of 2-chloro-DOPA in chemical defense and underwater adhesion are considered.

Journal of Adhesion

14272051

10.1242/JEB.068221

Fracture toughness of locust cuticle

SUMMARY Insect cuticle is one of the most common biological materials, yet very little is known about its mechanical properties. Many parts of the insect exoskeleton, such as the jumping legs of locusts, have to withstand high and repeated loading without failure. This paper presents the first measurements of fracture toughness for insect cuticle using a standard engineering approach. Our results show that the fracture toughness of cuticle in locust hind legs is 4.12 MPa m1/2 and decreases with desiccation of the cuticle. Stiffness and strength of the tibia cuticle were measured using buckling and cantilever bending and increased with desiccation. A combination of the cuticle’s high toughness with a relatively low stiffness of 3.05 GPa results in a work of fracture of 5.56 kJ m–2, which is amongst the highest of any biological material, giving the insect leg an exceptional ability to tolerate defects such as cracks and damage. Interestingly, insect cuticle achieves these unique properties without using reinforcement by a mineral phase, which is often found in other biological composite materials. These findings thus might inspire the development of new biomimetic composite materials.

The Journal of Experimental Biology

201659466

10.1002/ADMA.201903526

Avoidance of Catastrophic Structural Failure as an Evolutionary Constraint: Biomechanics of the Acorn Weevil Rostrum.

The acorn weevil (Curculio Linnaeus, 1758) rostrum (snout) exhibits remarkable flexibility and toughness derived from the microarchitecture of its exoskeleton. Modifications to the composite profile of the rostral cuticle that simultaneously enhance the flexibility and toughness of the distal portion of the snout are characterized. Using classical laminate plate theory, the effect of these modifications on the elastic behavior of the exoskeleton is estimated. It is shown that the tensile behavior of the rostrum across six Curculio species with high morphological variation correlates with changes in the relative layer thicknesses and orientation angles of layers in the exoskeleton. Accordingly, increased endocuticle thickness is strongly correlated with increased tensile strength. Rostrum stiffness is shown to be inversely correlated with work of fracture; thus allowing a highly curved rostrum to completely straighten without structural damage. Finally, exocuticle rich invaginations of the occipital sutures are identified both as a likely site of crack initiation in tensile failure and as a source of morphological constraint on the evolution of the rostrum in Curculio weevils. It is concluded that avoidance of catastrophic structural failure, as initiated in these sutures under tension, is the driving selective pressure in the evolution of the female Curculio rostrum.

Advanced Materials

406698

10.1098/RSPB.2009.2029

Induced niche shift as an anti-predator response for an endoparasitoid

When two developmental stages do not share the same ecological niche, the control of the niche shift through a change in developmental timing, referred to as ‘heterokairy’, can provide an adaptive advantage for the individual (e.g. if mortality risk is higher in the first niche). For endoparasitic species that develop inside another (host) species, mortality of the host may directly induce mortality risk for the parasite. Thus, endoparasitoid larvae should be selected for response to host predation. In this study, aphids previously parasitized by the endoparasitoid Endaphis fugitiva, Gagné and Muratori (Diptera: Cecidomyiidae), were experimentally exposed to increased mortality risks. Both simulated attack and actual predator attacks against aphid hosts induced early emergence of the parasitoid larvae. Parasitoid emergence from the aphids occurred several minutes before the predator finished feeding on the aphid, allowing enough time for the parasitoid larvae to avoid direct predation. Predator-induced emergence produced significantly smaller parasitoid larvae than controls, but, interestingly, no effect on Endaphis adult size was found. To our knowledge, this is the first evidence of induced emergence in an insect parasitoid, but we suggest that this mechanism might be at work in many other species where plasticity in development time allows the individual to perform an adaptive niche shift.

Proceedings of The Royal Society B: Biological Sciences

21333149

10.1093/AOB/MCS186

Collenchyma: a versatile mechanical tissue with dynamic cell walls.

BACKGROUND Collenchyma has remained in the shadow of commercially exploited mechanical tissues such as wood and fibres, and therefore has received little attention since it was first described. However, collenchyma is highly dynamic, especially compared with sclerenchyma. It is the main supporting tissue of growing organs with walls thickening during and after elongation. In older organs, collenchyma may become more rigid due to changes in cell wall composition or may undergo sclerification through lignification of newly deposited cell wall material. While much is known about the systematic and organographic distribution of collenchyma, there is rather less information regarding the molecular architecture and properties of its cell walls. SCOPE AND CONCLUSIONS This review summarizes several aspects that have not previously been extensively discussed including the origin of the term 'collenchyma' and the history of its typology. As the cell walls of collenchyma largely determine the dynamic characteristics of this tissue, I summarize the current state of knowledge regarding their structure and molecular composition. Unfortunately, to date, detailed studies specifically focusing on collenchyma cell walls have not been undertaken. However, generating a more detailed understanding of the structural and compositional modifications associated with the transition from plastic to elastic collenchyma cell wall properties is likely to provide significant insights into how specific configurations of cell wall polymers result in specific functional properties. This approach, focusing on architecture and functional properties, is likely to provide improved clarity on the controversial definition of collenchyma.

Annals of Botany