The climbing habit in palms: Biomechanics of the cirrus and flagellum

Climbing palms in the Arecoideae (Desmoncus) and Calamoideae (rattan palms) both evolved cirrate leaves armed with hooks and grapnels for climbing. Some species of Calamoideae develop a different climbing organ known as the flagellum, which also bears hooks. The present study indicates that geometry and mechanical properties of the cirrus vary between species. Cirrate leaves are constructed to optimize bending and torsion in relation to the deployment of recurved hooks. Hook development, size, and strength vary along cirri and flagella and are consistent with observations of these attachment organs functioning as a ratchet mechanism: hooks increase in strength toward the base of attachment organs and always fail before the axis in strength tests. Hook size and strength differ between species and are related to body size and ecological preference. Larger species produce larger hooks, but smaller climbing palms of the understory deploy fine sharp hooks that are effective on small diameter supports as well as large branches and trunks. The ephemeral nature of climbing organs in palms provides a challenge to their life-history development, particularly in terms of mechanical constraints and remaining attached to the host vegetation; these differ significantly from many vines and lianas having more perennial modes of attachment.     

Diversity of Mechanical Architectures in Climbing Plants: An Evolutionary Perspective

Mechanical architectures of a wide range of climbing plants are reviewed from a wide phylogenetic range and evolutionary contingencies. They include an herbaceous lycopod (Lycopodiaceae) - a woody tropical liana (Apocynaceae), temperate climbers, herbs and shrubs (Ranunculacae), and two representative climbing palms (Arecoideae, Calamoideae). Trends in mechanical properties during development are reviewed and interpreted via changes in anatomical development of the stem and type of connection to host supports. The results indicate that there are some biomechanical features common to diverse climbing plants including (1) phases of relatively rigid stem growth where the climbing stem has to span between supports and (2) a mechanism to achieve greater compliancy towards the base or at points where the slender climbing stem is at risk from excessive mechanical stress. Evolutionary contingencies such as basal plesiomorphic constraint, complexification, simplification and developmental loss can drastically influence ways in which different plants have evolved different biomechanical climbing architectures. Two key developmental features controlling the biomechanics of the climbing stem are (1) the presence/absence of secondary growth and (2) the number, complexity and coordination of development of primary or secondary tissues with varying mechanical properties. Recent research has suggested that evolution of specialized climbing architectures can canalize subsequent evolution of alternative growth forms. The results suggest that the origin and type of climbing architecture can be heavily influenced by ancestral growth forms and architectures. Despite the extremely complex patterns of plant growth form evolution involving escapes to and from more specialized or simpler bauplans, selective pressure towards non-self-supporting growth forms is a remarkably persistent and iterative feature of growth form evolution in land plants.     

Mechanical role of the leaf sheath in rattans

Leaf sheaths of rattans are long, tubular and persistent and unlike many self-supporting palms, extend far from the apex of the plant. The mechanical role of the leaf sheath was investigated in eight rattan species of the subfamily Calamoideae. The main objective was to analyse its influence on the mechanical architecture and contribution to the climbing habit. Bending mechanical properties were measured along climbing axes before and after removal of leaf sheaths. Results were related to stem and leaf sheath geometry and mechanical properties. Contribution of the leaf sheath to axial flexural rigidity was high (c. 90%) in the early stages of growth and towards the apex of older climbing axes for all climbing palms tested. Senescence and loss of the leaf sheath strongly influenced axial stiffness. A nonclimbing species, Calamus erectus, showed a different mechanical architecture. Although lacking secondary growth, palms have been able to develop successful climbers with a mechanical architecture broadly analogous to, although developmentally different from, dicotyledonous lianas. The role of the leaf sheath in modulating mechanical properties during ontogeny ought not to be neglected in studies on monocotyledons, as it possibly contributed significantly to the ways in which different growth forms have evolved in the group.     

Developmental plasticity and biomechanics of treelets and lianas in Manihot aff. quinquepartita (Euphorbiaceae): a branch-angle climber of French Guiana

Background and Aims

Most tropical lianas have specialized organs of attachment such as twining stems, hooks or tendrils but some do not. Many climbers also have an early self-supporting phase of growth and in some species this can produce treelet-sized individuals. This study focuses on how a liana can climb without specialized attachment organs and how biomechanical properties of the stem are modulated between self-supporting treelets and canopy-climbing lianas.

Methods

Biomechanics and stem development were investigated in self-supporting to climbing individuals of Manihot aff. quinquepartita (Euphorbiaceae) from tropical rain forest at Saül, central French Guiana. Bending tests were carried out close to the site of growth. Mechanical properties, including Young''s elastic modulus, were observed with reference to habit type and changes in stem anatomy during development.

Key Results

This liana species can show a remarkably long phase of self-supporting growth as treelets with stiff, juvenile wood characterizing the branches and main stem. During the early phase of climbing, stiff but unstable stem segments are loosely held in a vertical position to host plants via petiole bases. The stiffest stems – those having the highest values of Young''s modulus measured in bending – belonged to young, leaning and climbing stems. Only when climbing stems are securely anchored into the surrounding vegetation by a system of wide-angled branches, does the plant develop highly flexible stem properties. As in many specialized lianas, the change in stiffness is linked to the development of wood with numerous large vessels and thin-walled fibres.

Conclusions

Some angiosperms can develop highly effective climbing behaviour and specialized flexible stems without highly specialized organs of attachment. This is linked to a high degree of developmental plasticity in early stages of growth. Young individuals in either open or closed marginal forest conditions can grow as substantial treelets or as leaning/climbing plants, depending on the availability of host supports. The species of liana studied differs both in terms of development and biomechanics from many other lianas that climb via twining, tendrils or other specialized attachment organs.Key words: Biomechanics, bending, developmental plasticity, French Guiana, liana, Manihot aff. quinquepartita (Euphorbiaceae), treelet, branch angle climber, Young''s modulus     

Floristic diversity of sabal palmetto woodland: an endemic and endangered vegetation type from Mexico

The sabal palmetto woodland is a tropical plant formation dominated by Sabal mexicana, with restricted distribution to southeast Mexico. Sabal palms grow on poor soils but accumulate large quantities of organic substrate in their crowns, harboring a contingent of plants that use it as phorophyte. Although it is a threatened formation, basic information on its biodiversity is scant. We examined the floristic diversity of recruited (diameter at breast height, DBH, ≥1 cm) and understory (DBH ≤ 1 cm) plants, and its variation with anthropogenic disturbance. We also examined the floristic diversity of plants that use the sabal palms as phorophytes, and assessed its variation with human impact. All plants present in transects within a conserved and an adjacent perturbed area were sampled. The list of observed taxa shows that this vegetation has a clear affinity with tropical dry and wet forests of Mexico, with a small representation of taxa from desert ecosystems. The floristic contingent included a total of 81 species in 2000 m². Richness, composition and diversity were affected by disturbance. Recruited and understory vegetation in the disturbed site were 5- and 1.6-times less diverse than in the conserved site, and species of mature, conserved vegetation were substituted by heliophytes in the disturbed site. In contrast, abundance of palms and diversity and identity of epiphytic/hemiepiphytic plants were not affected by disturbance. We show that even monodominated tropical ecosystems growing on poor soils have a high floristic diversity and that current anthropogenic impact threatens not only species and populations but also entire plant formations.     

Structural rigidity of paranemic crossover and juxtapose DNA nanostructures

Crossover motifs are integral components for designing DNA-based nanostructures and nanomechanical devices due to their enhanced rigidity compared to the normal B-DNA. Although the structural rigidity of the double helix B-DNA has been investigated extensively using both experimental and theoretical tools, to date there is no quantitative information about structural rigidity and the mechanical strength of parallel crossover DNA motifs. We have used fully atomistic molecular dynamics simulations in explicit solvent to get the force-extension curve of parallel DNA nanostructures to characterize their mechanical rigidity. In the presence of monovalent Na⁺ ions, we find that the stretch modulus (γ₁) of the paranemic crossover and its topoisomer JX DNA structure is significantly higher (∼30%) compared to normal B-DNA of the same sequence and length. However, this is in contrast to the original expectation that these motifs are almost twice as rigid compared to the double-stranded B-DNA. When the DNA motif is surrounded by a solvent with Mg²⁺ counterions, we find an enhanced rigidity compared to Na⁺ environment due to the electrostatic screening effects arising from the divalent nature of Mg²⁺ ions. To our knowledge, this is the first direct determination of the mechanical strength of these crossover motifs, which can be useful for the design of suitable DNA for DNA-based nanostructures and nanomechanical devices with improved structural rigidity.     

Disruption of recruitment in two endemic palms on Lord Howe Island by invasive rats

Invasive species may have negative impacts on many narrow range endemics and species restricted to oceanic islands. Predicting recent impacts of invasive species on long-lived trees is difficult because the presence of adult plants may mask population changes. We examined the impact of introduced black rats (Rattus rattus) on two palm species restricted to cloud forests and endemic to Lord Howe Island, a small oceanic island in the southern Pacific. We combined estimates of the standing size distribution of these palms with the proximal impacts of rats on fruit survival in areas baited to control rats and in unbaited areas. The size distribution of palms with trunks was comparable across baited and unbaited sites. Small juvenile palms lacking a trunk (<50 cm tall) were abundant in baited areas, but rare in unbaited sites for Lepidorrhachis mooreana, and rare or absent in 3 out of 4 unbaited Hedyscepe canterburyana sites. All ripe fruits were lost to rats in the small fruited L. mooreana. Fruit removal was widespread but less (20–54%) in H. canterburyana. Both palms showed evidence of a reduced capacity to maintain a juvenile bank of palms through regular recruitment as a consequence of over 90 years of rat impact. This will limit the ability of these species to take advantage of episodic canopy gaps. Baiting for rat control reduced fruit losses and resulted in the re-establishment of a juvenile palm bank. Conservation of both endemic palms necessitates control (or eradication) of rat populations on the unique cloud forest summits of the island.     

LEAF DEVELOPMENT AND CROWN GEOMETRY OF TWO IRIARTEOID PALMS

We examined changes in pinnately compound leaf morphology and crown geometry that occur during height growth of the iriarteoid palms Socratea exorrhiza and Iriartea deltoidea in tropical wet forest of Costa Rica. Although light availability increased with height, the number of leaves per plant was relatively constant. Total leaf area, however, was much larger in taller individuals. Increases in linear dimensions of leaves (length and width) was responsible for less than half of this greater surface area. More important was the transition from a basically dorsiventral display of leaflets in small individuals to a more radial display in taller plants. Production of leaflets in more than one plane resulted in leaves whose surface area was more than twice the horizontally projected area and whose lateral light interception was greatly enhanced. S. exorrhiza, a faster-growing and more light-demanding species, undergoes this transformation in leaf morphology at heights between 3 and 6 m; whereas in the slower-growing and more shade-tolerant I. deltoidea this occurs at heights between 10 and 20 m. Pinnate leaves, with dense radial packing of leaflets along the rachis, are functionally comparable to branches of dicotyledonous trees and may have been important for the evolution of arborescence in palms.     

On Heels and Toes: How Ants Climb with Adhesive Pads and Tarsal Friction Hair Arrays

Ants are able to climb effortlessly on vertical and inverted smooth surfaces. When climbing, their feet touch the substrate not only with their pretarsal adhesive pads but also with dense arrays of fine hairs on the ventral side of the 3^rd and 4^th tarsal segments. To understand what role these different attachment structures play during locomotion, we analysed leg kinematics and recorded single-leg ground reaction forces in Weaver ants (Oecophylla smaragdina) climbing vertically on a smooth glass substrate. We found that the ants engaged different attachment structures depending on whether their feet were above or below their Centre of Mass (CoM). Legs above the CoM pulled and engaged the arolia (‘toes’), whereas legs below the CoM pushed with the 3^rd and 4^th tarsomeres (‘heels’) in surface contact. Legs above the CoM carried a significantly larger proportion of the body weight than legs below the CoM. Force measurements on individual ant tarsi showed that friction increased with normal load as a result of the bending and increasing side contact of the tarsal hairs. On a rough sandpaper substrate, the tarsal hairs generated higher friction forces in the pushing than in the pulling direction, whereas the reverse effect was found on the smooth substrate. When the tarsal hairs were pushed, buckling was observed for forces exceeding the shear forces found in climbing ants. Adhesion forces were small but not negligible, and higher on the smooth substrate. Our results indicate that the dense tarsal hair arrays produce friction forces when pressed against the substrate, and help the ants to push outwards during horizontal and vertical walking.     

Seasonal variation in agar composition and properties from Gracilaria gracilis (Gracilariales,Rhodophyta) of the Patagonian coast of Argentina

Seasonal variation of agar from specimens of a commercially exploited population of Gracilaria gracilis (Stackhouse) Steentoft, Irvine & Farnham in the Patagonian coast of Argentina was studied. For each seasonal harvest, random samples of plants were pooled for subsequent polysaccharide extraction at different water temperatures and agar physico‐chemical properties and composition were determined. Both spring and summer plants yielded 30% and 41% of agar, respectively, which differed slightly in their at rest rheological behavior. Spring and summer plants produced strong gels (238 and 218 g cm^?2, respectively), but the latter gels had nil adhesiveness. In autumn plants, agar yield decayed to 19%, though the product still maintained similar gel strength (210 g cm^?2). Adhesiveness in this product was at least an order of magnitude higher than in the others, concomitant with a peak in the formation of tetraspores and carpospores. This suggests a biological role for the galactan in the initial attachment of spores to the substrate. But since fall corresponds to the settling of reproductive structures, caution should be taken to harvest the algae once spores have been shed from mother plants.     

Portamento Rampicante - Volubile Provocato,in Alcune Piante,dal Trattamento Con Acido Gibberellico

Summary

In this communication reports are given on experiments and observations regarding treatment of several plants with Gibberellic acid and consequent manifestations of stem growth with twining-climbing behaviour. This phenomenon has been observed in plants which normally never present similar behaviour (Lactuca Scariola L.) and it has been provoked in plants genetically more or less predisposed (Phaseolus vulgaris L. cult. “aquila rossa” e Soja hispida Moench. cult. “Castelli” and “Giessner”). Normally climbing plants, as Ipomaea versicolor, has been treated for control.

In Phaseolus and in Soja the action of Gibberellic acid has been very considerable as plants thus treated have assumed all the characteristics of climbing growth. Photos and Tables clearly show the extent of such action.

In Ipomaea, in conformity with work hypothesis, we can clearly observe the meagre advantage in growth of internodes following tratment with Gibberellic Acid.

The purpose of these experiments, as is obvious, is to lay the bases for a physiological interpretation of climbing behaviour. It is clear by results obtained that, subordinately, at least, to certain qualitative structural manifestations, plants thrive with or without climbing behaviour according to whether their growth is more or less rapid and of different proportions. Increasing growth rapidity by tratment with Gibberellic Acid, we may obtain an twining growth phenomenon in plants genetically more or less predisposed towards said phenomenon, in those plants, that is, which in special environnmental conditions may sometimes, although exceptionally, manifest the tendency to climbing growth. This effect is obtained on plants in which we do not usually meet these phenomena but which, evidently, have the possibility of manifest them provided that am exceptional growth stimulas is operating.

From what above said we can deduce a first and simple work hypothesis, which will be the object of study in the near future, and which prospects the possibility that typically climbing plants owe their behaviour to the capacity of regulating their growth also in consequence of the formation of appropriate quantities of substances with gibberellinlike action and lacking some growth inhibiting condition. The same thing might be true for the plants which are climbing only on exceptional occasions, and a similar possibility might be given to the plant by particular external conditions in which the individual is formed, grows and develops.

Is is interesting also to observe that the analogy between climbing behaviour induced by Gibberellic Acid and natural one, extends also to leaf expansion behaviour and to other characteristics facts included in the field of etiolated plants.

Successive experiments will give us a deeper knowledge of this particular aspect of climbing behaviour and will define the conditions in which movements of shoot apex take place under the action of Gibberellic Acid and structural manifestations inherent, in comparison to those of naturally climbing plants.     

Dry-matter production and nutrient content of plantation oil palms in Nigeria

Summary Seventeen palms selected as representative of six age-groups ranging from 7 to 22 years were cut down, split into their component parts and the dry weights obtained. Data on plant composition and dry-matter accumulation since planting are given together with estimates of annual production expressed as dry matter per hectare (crop-growth rate) and on the basis of leaf area (net assimilation rate). The results are discussed in relation to productivity of other plants in temperate regions and in the tropics, and reasons are suggested for the comparatively low rate of dry-matter production (1.95 × 10⁴ kg/ha/annum) in adult palms.     

Structure, attachment properties, and ecological importance of the attachment system of English ivy (Hedera helix)

Root climbers such as English ivy (Hedera helix) rely on specialized adventitious roots for attachment, enabling the plants to climb on a wide range of natural and artificial substrates. Despite their importance for the climbing habit, the biomechanical properties of these specialized adventitious roots compared with standard roots and their performance in the attachment to different host species or inert substrates have not been studied. Here organs and tissues involved in the attachment are characterized and their significance in regard to a broader functional and ecological aspect is discussed. Depending on the substrate, the root clusters show different types of failure modes at various frequencies, demonstrating the close interaction between the climber and its substrates. With a Young's Modulus of 109.2 MPa, the attachment roots are relatively stiff for non-woody roots. The central cylinders of the attachment roots show a high tensile strength of 38 MPa and a very high extensibility of 34%. In host trees naturally co-distributed with English ivy, a 'balanced' occurrence of failure of the attachment system of the climber and the bark of the host is found, suggesting a co-evolution of climber and host. Maximum loads of root clusters normalized by the number of roots match those of individually tested attachment roots. In comparison with most subterranean roots the properties and structure of the attachment roots of English ivy show distinct differences. There exist similarities to the properties found for roots of Galium aparine, suggesting a trend in not fully self-supporting plants towards a higher extensibility.     

薇甘菊不同生长方式下的繁殖特征比较

利用实验生态学方法, 对攀缘生长与伏地生长的薇甘菊(Mikania micrantha)的形态特征、生物量分配和繁殖特征进行了比较研究。结果表明: 在潮湿生境条件下, 攀缘生长的植株与伏地生长的植株相比对主茎的生物量投资较大, 而对分枝茎的生物量投资则相对较小; 但在干旱生境条件下, 攀缘生长的植株对主茎和分枝茎的生物量投资均较伏地生长的植株小。在潮湿与干旱生境中, 攀缘植株的繁殖投资分别为0.429 g·g^–1和0.342 g·g^–1, 显著高于相同生境中的伏地植株。在薇甘菊种群的补充与更新过程中, 攀缘生长的植株以有性生殖占主导地位, 在潮湿与干旱生境中产生的个体分别占种群数量的84.7%和62.6%; 伏地生长的植株则主要以无性(克隆)繁殖为主, 在潮湿与干旱生境中有性生殖产生的个体仅占种群数量的40.4%和35.9%。生长方式与土壤水分二因子互作效应对薇甘菊的生物量分配和繁殖指标影响均达到显著水平(P<0.05)。通过研究得出以下结论: 不同生长方式的薇甘菊对环境条件具有不同的生长与繁殖适应对策。     

Water resource partitioning,stem xylem hydraulic properties,and plant water use strategies in a seasonally dry riparian tropical rainforest

This study investigated seasonal variation in the origin of water used by plants in a riparian tropical rainforest community and explored linkages between plant water source, plant xylem hydraulic conductivity and response to the onset of dry conditions. The study focused on five co-dominant canopy species, comprising three tree species (Doryphora aromatica, Argyrodendron trifoliolatum, Castanospora alphandii) and two climbing palms (Calamus australis and Calamus caryotoides). Stable isotope ratios of oxygen in water (¹⁸O) from soil, groundwater, stream water and plant xylem measured in the wet season and the subsequent dry season revealed water resource partitioning between species in the dry season. Measurement of stem-area-specific hydraulic conductivity (K_S) in the wet season and subsequent dry season showed a significant dry-season loss of K_S in three of the five species (Castanospora alphandii, Calamus australis and C. caryotoides) and a decrease in mean K_S for all species. This loss of hydraulic conductivity was positively correlated with the difference between wet-season and dry-season midday leaf water potentials and with leaf carbon isotope discrimination, indicating that plants that were less susceptible to loss of conductivity had greater control over transpiration rate and were more water-use efficient.     

Biomechanics and development of the climbing habit in two species of the South American palm genus Desmoncus (Arecaceae)

Mechanical properties are investigated in Desmoncus orthacanthos and D. polyacanthos from French Guiana, South America. Differences in size and axis stiffness are related to different trellis requirements and habitats. The leaf sheath surrounds the stem, increasing stiffness of young self-supporting stages and apical parts of older climbing plants. Senescence of the leaf sheath reduces stiffness of older climbing axes of both species. Its eventual loss in D. orthacanthos facilitates deformation into coils and loops when plants slip from their supports following senescence of leaves bearing attachment organs. In smaller climbing axes of D. polyacanthos, the senescent leaf sheath remains attached and axes rarely form loops and coils below attachment. An increase in stiff mechanical properties toward the base of both species is radically different from that of many dicotyledonous lianas. Besides the presence of attachment organs, stem mechanical properties of Desmoncus are similar to those of erect though not fully self-supporting stems of Bactris major, a sympatric species of the sister group genus to Desmoncus. The climbing habit in Desmoncus may have evolved via (1) heterochronic processes including early elongation of internodes relative to increase in stem diameter (reduction of the establishment phase) and (2) increased persistence of leaf sheaths.     

Establishment growth of cabbage palm,sabal palmetto (arecaceae)

Cabbage palms (Sabal palmetto) go through an establishment phase during which the stem grows downward for a period of time before growing upward. We estimated the duration of this phase for cabbage palms growing in coastal forest in Florida using a matrix model approach. All data were collected over a 2-yr period (1993-1995) in coastal forest at Waccasassa∗∗∗ Bay, Florida. The minimum time projected by the model for a plant to develop an aboveground trunk was 14 yr. We estimate that the fastest growing 1, 10, and 50% of plants would develop an aboveground trunk in 33, 42, and 59 yr, respectively. The projected duration of the trunkless phase is surprisingly long but not unlike other palms with similar types of establishment growth. Our estimates are much longer than anecdotal estimates for cabbage palms grown under nursery conditions but are similar to anecdotal estimates for plants grown in field conditions. Management practices that remove cabbage palms with aboveground stems usually leave a population of palms without aboveground stems that serves as temporary reserve for relatively rapid recovery of the palm stand. This may foster the perception by many that cabbage palms are fast growing.     

Ecological applications of differences in the hydraulic efficiency of palms and broad-leaved trees

Key message

In tropical forests, co-occurring woody monocot and dicot species adapted different water use strategies highly depending on their investment in the hydraulic conduit properties.

Abstract

We studied the hydraulic efficiency of palms and broad-leaved tropical tree species from a moist tropical lowland forest in the Central Brazilian Amazon. Therefore, we harvested 34 trees and 10 açai palms and measured vessel size and frequency at diameter at breast height and additionally at the base of the crown shaft for the palms. Further, we assessed the active xylem area to estimate the hydraulic conductivity through Hagen Poiseuille’s adapted theoretical equation. Mean vessel diameter in dicot trees was 127.62 ± 49.22 μm with an average 9.09 ± 6.50 vessels per mm². Mean conduits sizes at the base (h = 0.10 m) of palm trees were larger with 288.20 ± 32.96 μm and less frequent with 1.40 ± 0.46 vessels per mm². Hydraulic conductivity was on average 3.31 ± 4.59 kg m^?1 s^?1 MPa^?1 for dicot trees. Mean hydraulic conductivity in açai palms was 20.45 ± 10.6 kg m^?1 s^?1 MPa^?1 at the base, and increased to 124.73 ± 55.2 kg m^?1 s^?1 MPa^?1 at the crown base. Hydraulic conductivity at the base of the crown was higher than in the base of the trunk due to the high density of vessels in a small cross-section in this height. Furthermore, we found a species-independent relationship between vessel diameter and frequency. We conclude that the differences found in the hydraulic efficiency give some evidence that palms have a lower occurrence of embolism and cavitation than trees, which is due to stiffer and stronger conduit pathways and efficient drought-avoiding strategies. The differences in hydraulic architecture between palms and trees imply different water use patterns thus varying niche differentiation, but this does not consequently need to be an excluding factor for coexistence in the same environment.

     

Corytibacladius Oliveira,Messias & Santos, 1995, a junior synonym of Limnophyes Eaton, 1875 (Diptera: Chironomidae: Orthocladiinae)

Larvae of the blackfly Simulium noelleri Friederichs aggregate at high populatio n densities on dams and spillways at the outlet of ponds. When displaced into the water column from their point of attachment, larvae can secrete silk threads as "life-lines" which enable them to maintain a link to the substratum an d up which they can climb to regain their original position. Experiments were conducted in a laboratory pond outlet to investigate this use of silk threads, larvae being displaced by means of forceps. It was demonstrated that: (i) the length of thread produced , and the speed of climbing the thread are independent of larval size; (ii) within limits, the speed of climbing was independent of both the length of the thread and the time already spent climbing, and (iii) speed of climb became more rapid as larvae neared the point of attachment. The range of locomotion in blackfly larvae is then discussed.     

Geometry and bone mineral density determinants of femoral neck strength changes following exercise

Physical exercise induces spatially heterogeneous adaptation in bone. However, it remains unclear where the changes in BMD and geometry have the greatest impact on femoral neck strength. The aim of this study was to determine the principal BMD-and-geometry changes induced by exercise that have the greatest effect on femoral neck strength. Pre- and post-exercise 3D-DXA images of the proximal femur were collected of male participants from the LIFTMOR-M exercise intervention trial. Meshes with element-by-element correspondence were generated by morphing a template mesh to each bone to calculate changes in BMD and geometry. Finite element (FE) models predicted femoral neck strength changes under single-leg stance and sideways fall load. Partial least squares regression (PLSR) models were developed with BMD-only, geometry-only, and BMD-and-geometry changes to determine the principal modes that explained the greatest variation in neck strength changes. The PLSR models explained over 90% of the strength variation with 3 PLS components using BMD-only (R² > 0.92, RMSE < 0.06 N) and 8 PLS components with geometry-only (R² > 0.93, RMSE < 0.06 N). Changes in the superior neck and distal cortex were most important during single-leg stance while the superior neck, medial head, and lateral trochanter were most important during a sideways fall. Local changes in femoral neck and head geometry could differentiate the exercise groups from the control group. Exercise interventions may target BMD changes in the superior neck, inferior neck, and greater trochanter for improved femoral neck strength in single-leg stance and sideways fall.