Intra-annual wood anatomical features of high-elevation conifers in the Great Basin,USA

Mountain conifers in the Great Basin of North America have provided some of the longest, continuous, and annually resolved paleoclimate records. Climate-growth relationships at the cellular level, which help understand wood formation processes that underlie dendroclimatic reconstructions, are at present largely unexplored in the Great Basin. We analyzed 42 trees located in the Snake Range (eastern Nevada, USA) at three sites along an elevation gradient. Sampled species included white fir (Abies concolor), Douglas fir (Pseudotsuga menziesii), limber pine (Pinus flexilis), bristlecone pine (Pinus longaeva), and Engelmann spruce (Picea engelmannii). Wood anatomical features were quantified for two consecutive years, 2011 and 2012. Lumen area, cell wall thickness, lumen diameter, and wall-to-cell ratio were measured for the total ring as well as for earlywood and latewood. Mean standardized tracheidograms highlighted differences between 2011 and 2012, in particular concerning lumen area and wall-to-cell ratio. Most annual variation was due to earlywood, rather than latewood. Anatomical parameters of limber pine, the only species that could be tested at both the montane and subalpine sites, varied with elevation. Principal component analysis showed that the main axis of variability was related to dimensional parameters (e.g. lumen area), which reflected differences in water availability.     

Morphology and ecological significance of intra-annual radial cracks in living conifers

 Intra-annual radial cracks were studied on 294 cross-sections of Norway spruce sampled at two forest sites in the eastern Alps (Italy) and from seven isolated trees in the Jura region (Switzerland). Cracks were occasionally accompanied by traumatic resin canals in the wood that was formed after the cracking. Most of the cracks, however, were without such canals. Traumatic resin canals are not significantly more abundant in tree rings formed after cracking, and their occurrence is not related to the cracking. Cracks developed when the cambium was inactive. Water imbalances during the early spring, due to transpiration losses and inadequate moisture supply from very cold roots, are the likely cause of these cracks. Received: 21 February 1996 / Accepted: 14 June 1996     

Adjusting xylem anatomy and growth to inter-annual climate variability in two Fabaceae species (Centrolobium microchaete,Cenostigma pluviosum) from Bolivian dry tropical forests

Climate-growth relationships are strong in seasonally dry tropical forests (SDTFs). To better understand the ecological processes controlling these relationships we need to assess the long-term responses of wood anatomy and radial growth to year-to-year climate variability. We assessed how wood-anatomical traits (mean vessel area –MVA– and vessel density –VD–, percentage of conductive area –CA–, xylem-specific estimated hydraulic conductivity –Ks–) and growth responded to local climate (mean temperature, total precipitation, estimated moisture) variability and teleconnections (Pacific Decadal Oscillation) between 1970 and 2011 in two Fabaceae tree species (Centrolobium microchaete and Cenostigma pluviosum) coexisting in a Bolivian SDTF. We found that C. microchaete produced wider vessels and was more responsive to both local climate conditions and teleconnections than C. pluviosum. In C. microchaete VD positively responded to average temperature in the late-wet season and in the previous dry season, and CA and Ks were higher in years with warmer wet and previous early-dry seasons, as well as in years with higher PDO values. These responses were independent from ring-width variability only for C. microchaete. For C. pluviosum, vessel chronologies were more responsive to local temperature variability, and only MVA and CA positively and negatively responded to moisture in the early-dry and the previous dry seasons, respectively. Our results show that wood hydraulic structure in SDTFs is responsive to climate fluctuations. The combined study of ring width and wood anatomy allows having a more complete picture of the influence of climate on growth, particularly in species as C. microchaete which show low collinearity of tree-ring width and wood plasticity in response to climate variability.     

Studying global change through investigation of the plastic responses of xylem anatomy in tree rings

Variability in xylem anatomy is of interest to plant scientists because of the role water transport plays in plant performance and survival. Insights into plant adjustments to changing environmental conditions have mainly been obtained through structural and functional comparative studies between taxa or within taxa on contrasting sites or along environmental gradients. Yet, a gap exists regarding the study of hydraulic adjustments in response to environmental changes over the lifetimes of plants. In trees, dated tree-ring series are often exploited to reconstruct dynamics in ecological conditions, and recent work in which wood-anatomical variables have been used in dendrochronology has produced promising results. Environmental signals identified in water-conducting cells carry novel information reflecting changes in regional conditions and are mostly related to short, sub-annual intervals. Although the idea of investigating environmental signals through wood anatomical time series goes back to the 1960s, it is only recently that low-cost computerized image-analysis systems have enabled increased scientific output in this field. We believe that the study of tree-ring anatomy is emerging as a promising approach in tree biology and climate change research, particularly if complemented by physiological and ecological studies. This contribution presents the rationale, the potential, and the methodological challenges of this innovative approach.     

Neotropical Hypericum irazuense shrubs reveal recent ENSO variability in Costa Rican páramo

Climate-vegetation relations in alpine systems play a pivotal role in regulating hydrology and have thus become a research priority in a context of ongoing climate change. In this paper, we investigate how one of the most dominant shrub species in alpine páramo ecosystems of Central America, Hypericum irazuense, responds to changes in precipitation, temperature and El Niño-Southern Oscillation. To this end, we performed dendrochronological and wood-anatomical analyses on H. irazuense to determine the limiting climatic factors driving shrub growth, using a bootstrapped correlation and response function analysis. To validate our results further, we also applied Structural Equation Models (SEM), an approach commonly used in ecology, so as to check for climate-growth relations which consider the control of ENSO on growth through its influence on various climatic parameters. Results support a relation between climate and annual growth of H. irazuense and demonstrate that the latter is sensitive to precipitation and temperature during boreal winters. In addition, we observe a statistically significant correlation between annual growth and La Niña events. The presence of annual growth rings holds H. irazuense as one in only few neotropical species suited for dendrochronological studies. Results of this study could thus contribute to an improved understanding of how changing climatic conditions affect the fragile and threatened páramo ecosystem and the ensuing services it offers in the form of hydrology regulation over the next decades.     

X-ray microdensitometry applied to subfossil tree-rings: growth characteristics of ancient pines from the southern boreal forest zone in Finland at intra-annual to centennial time-scales

A collection of subfossil wood of Pinus sylvestris (Scots pine) was exposed to X-ray densitometry. The collection of 64 samples from the southern boreal forest zone was dendrochronologically cross-dated to a.d. 673-1788. Growth characteristics were determined by performing density profiles including the following parameters: minimum density, earlywood and latewood boundary density, maximum density, earlywood width, earlywood density, latewood width, latewood density, annual ring width and annual ring density. Seven out of the nine parameters were found to contain non-climatic growth trends and six were found to be heteroscedastic in their variance. Tree-specific records were indexed, to remove the non-climatic growth trends and stabilize the variance, and combined into nine parameter-specific tree-ring chronologies. Growth characteristics of the pines changed in parallel with the generally agreed climatic cooling from the Medieval Warm Period to the Little Ice Age: pine tree-rings showed decreasing maximum densities from the period a.d. 975-1150 to a.d. 1450–1625. A concomitant change in the intra-annual growth characteristics was detected between these periods. The findings indicate that not only the trees growing near the species’ distributional limits are sensitive to large-scale climatic variations but also the trees growing in habitats remote from the timberline have noticeably responded to past climate changes.     

青藏高原高山植物的形态和解剖结构及其对环境的适应性研究进展

高山植物是一类生长于树线以上至雪线的山地植物。揭示高山植物适应环境的形态和结构特征及其内在机制,对研究全球气候变化下,植物对环境的响应和适应具有重要的理论意义。然而,国内在高山植物功能生态学的研究上相对薄弱,已有研究主要集中在对青藏高原高山植物的报道上。结合国外高山植物的相关研究报道,从形态和解剖结构两个方面对青藏高原高山植物的研究进展进行了综述,重点阐述了高山植物的形态、解剖结构及其与环境的适应性关系。植株矮小（有的呈垫状）、叶片小而厚、具有通气组织、栅栏组织多层、机械组织发达、虫媒花性状、线粒体数量多和叶绿体基粒片层少等是这一地区高山植物普遍具有的形态和结构特征。高山植物形成上述结构的特异性是高山特殊综合生态环境长期作用的结果。同时,也是高山植物对高山环境的高度适应。最后,对这一领域存在的问题以及未来研究的重点和方向进行了探讨。目的是引起国内研究者的关注,促进我国高山植物功能生态学的研究与发展。     

A latent variable approach to study gene-environment interactions in the presence of multiple correlated exposures

Many existing cohort studies initially designed to investigate disease risk as a function of environmental exposures have collected genomic data in recent years with the objective of testing for gene-environment interaction (G × E) effects. In environmental epidemiology, interest in G × E arises primarily after a significant effect of the environmental exposure has been documented. Cohort studies often collect rich exposure data; as a result, assessing G × E effects in the presence of multiple exposure markers further increases the burden of multiple testing, an issue already present in both genetic and environment health studies. Latent variable (LV) models have been used in environmental epidemiology to reduce dimensionality of the exposure data, gain power by reducing multiplicity issues via condensing exposure data, and avoid collinearity problems due to presence of multiple correlated exposures. We extend the LV framework to characterize gene-environment interaction in presence of multiple correlated exposures and genotype categories. Further, similar to what has been done in case-control G × E studies, we use the assumption of gene-environment (G-E) independence to boost the power of tests for interaction. The consequences of making this assumption, or the issue of how to explicitly model G-E association has not been previously investigated in LV models. We postulate a hierarchy of assumptions about the LV model regarding the different forms of G-E dependence and show that making such assumptions may influence inferential results on the G, E, and G × E parameters. We implement a class of shrinkage estimators to data adaptively trade-off between the most restrictive to most flexible form of G-E dependence assumption and note that such class of compromise estimators can serve as a benchmark of model adequacy in LV models. We demonstrate the methods with an example from the Early Life Exposures in Mexico City to Neuro-Toxicants Study of lead exposure, iron metabolism genes, and birth weight.     

Wood anatomy and growth rate of seasonally dry tropical forest trees in the Marañón River Valley,northern Peru

Seasonally dry tropical forests (SDTF) currently occupy only 10% of the original area of their natural distribution. In the Marañón river valley, north western of Peru, occurs one of the most important SDTF of the neotropics. It has an endemic woody flora that represent 32% of the species, being threatened by its increasing fragmentation, and the knowledge of the ecology, forestry and conservation of the tree species is scarce. In this context this work presents the result of evaluation of the wood anatomy with emphasis to growth ring analysis, dating trees and stem growth rate determination of four tree species: Anadenanthera colubrina (Vell.) Brenan, Cedrela kuelapensis T.D. Penn. & Daza, Cordia iguaguana Melch. Ex I.M. Johnst., and Esenbeckia cornuta Eng., the last three being endemic and threatened with extinction. Our results determined that Cedrela kuelapensis showed the highest radial growth of the stem in relation to the other congeners of the neotropical region. Lower stem growth was observed for E. cornuta trees, according to the pattern of other tropical species. The results also indicate that populations of C. iguaguana and Esenbeckia cornuta in the zone are mainly composed by young trees exhibiting strong anthropic pressure due to the quality of their timber and, in consequence, more efforts are needed in the management of these resources to guarantee their sustainability.     

Vascular occlusion in stems of cut rose flowers exposed to air: Role of xylem anatomy and rates of transpiration

Stems of cut rose flowers were exposed to air at 20°C and 60% relative humidity and then placed in water. The rate of water uptake in Frisco, Sonia, Madelon, and Cara Mia roses was maximally inhibited after 72, 36, 24, and 3 h of exposure to air. respectively. Anatomical observations showed no tyloses, gums, or deposition of hydrophobic material in the xylem conduits (tracheids and vessels) of any of the investigated cultivars. Frisco, Sonia, Madelon, and Cara Mia roses showed no difference in the number, the length, the wall thickness, or the diameter of the lumina of either the tracheids or the vessels. This indicates that differences in recovery are not related to capillary diameter or length.
During exposure to air the decrease in transpiration rate, fresh weight, and water potential was the same in Sonia, Madelon, and Cara Mia roses. Upon exposure to air Frisco roses lost less water than the other cultivars studied In Frisco roses stomatal conductance was similar to that of other cultivars, but the rate of cuticular transpiration was lower.
It is concluded that conservation of water through low cuticular transpiration is one of the reasons for the relatively slow development of the vascular occlusion in Frisco roses, but the differences between Sonia. Madelon and Cara Mia roses were not related to their rates of transpiration.     

Applying climwin to dendrochronology: A breakthrough in the analyses of tree responses to environmental variability

Observational, correlative approaches are one of the backbones of dendrochronology. For instance, climate-growth relationships are usually quantified by calculating Pearson correlations. However, the ability to detect these relationships and the probability of declaring significant correlations by chance pose multiple challenges to such correlative framework. The R climwin package, developed a few years ago within the discipline of animal ecology, overcomes these limitations. In this paper we apply climwin to study relationships between climate and tree-ring widths and anatomy to show the advantages of using this package in the field of dendrochronology. This package allows calculating several models considering multiple windows relating a response variable to the climatic factors at different time resolutions. Then, the most parsimonious model is selected through an information-theoretic approach and randomization tests are computed to establish the significance of the selected model. We compare analyses based on Pearson correlations with climwin results using several environmental drivers (climate variables, drought indices, river flow), response variables (tree-ring width, tracheid lumen area and cell-wall thickness), and tree species from ecologically contrasting sites (cold- and water-limited conifers, Mediterranean riparian ash forests). Analyses of climate-growth/anatomy relationships based on the use of climwin showed several advantages over simple Pearson correlations: (i) they did not depend on the use of arbitrary time intervals of fixed duration, (ii) they allowed reducing probabilities associated with type I and II errors, (iii) they resulted in more consistent findings, (iv) they increased the capacity to detect differences between sites or periods in a time series, and (v) they provided more explanatory power.     

A theoretical approach to cluster formation in biological membranes

A theoretical analysis of cluster formation within the lipid matrix of biological membranes is presented. Various models are analysed: (a) one-dimensional monolayer, (b) two-dimensional monolayer and (c) one dimensional bilayer. Furthermore, lipid-protein interactions are considered. The model is based on differential equations for the probabilities a_i and b₁ which characterize the occupation of the lattice site i by the lipids A and B, respectively. These differential equations are an approximation of the Master-equation. Steady states as well as time-dependent variations are analysed. Depending on the interaction energies of the two lipids, different stationary lipid distributions are obtained, including clusters of lipids A or B and alternating structures. The distributions may be dynamically stable or unstable. It is shown that phase transitions within the lipid matrix may be induced by alteration of the composition of the membrane, by changing the interaction energies of the lipids, by variation of the temperature or by lipid-protein interactions. The transitions between different stationary distributions are studied by use of bifurcation diagrams. The analysis of time-dependent states reveals that unstable structures of the membrane may be important for certain time periods. Consideration of the lipid bilayer leads to a great number of possible distributions, which may be symmetric or asymmetric with respect to the outer and inner leaflets of the membrane.     

A functional trait approach to identifying life history patterns in stochastic environments

Temporal variation in demographic processes can greatly impact population dynamics. Perturbations of statistical coefficients that describe demographic rates within matrix models have, for example, revealed that stochastic population growth rates (log(λ_s)) of fast life histories are more sensitive to temporal autocorrelation of environmental conditions than those of slow life histories. Yet, we know little about the mechanisms that drive such patterns. Here, we used a mechanistic, functional trait approach to examine the functional pathways by which a typical fast life history species, the macrodetrivore Orchestia gammarellus, and a typical slow life history species, the reef manta ray Manta alfredi, differ in their sensitivity to environmental autocorrelation if (a) growth and reproduction are described mechanistically by functional traits that adhere to the principle of energy conservation, and if (b) demographic variation is determined by temporal autocorrelation in food conditions. Opposite to previous findings, we found that O. gammarellus log(λ_s) was most sensitive to the frequency of good food conditions, likely because reproduction traits, which directly impact population growth, were most influential to log(λ_s). Manta alfredi log(λ_s) was instead most sensitive to temporal autocorrelation, likely because growth parameters, which impact population growth indirectly, were most influential to log(λ_s). This differential sensitivity to functional traits likely also explains why we found that O. gammarellus mean body size decreased (due to increased reproduction) but M. alfredi mean body size increased (due to increased individual growth) as food conditions became more favorable. Increasing demographic stochasticity under constant food conditions decreased O. gammarellus mean body size and increased log(λ_s) due to increased reproduction, whereas M. alfredi mean body and log(λ_s) decreased, likely due to decreased individual growth. Our findings signify the importance of integrating functional traits into demographic models as this provides mechanistic understanding of how environmental and demographic stochasticity affects population dynamics in stochastic environments.     

From modeling the kinetics of cell enlargement to building tracheidograms of Larix gmelinii Rupr. (Rupr.) in the permafrost zone in Siberia

Boreal forest ecosystems are a crucial element in the global climate balance. In harsh environments functioning lateral meristems of trees are more regulated by the exogenous (included local climate) than endogenous factors. This information is encoded in the tree-ring structure which can be effectively decomposed by the process-based tree-ring growth simulations. Moreover, the process-based modeling can be used to describe non-linear processes linking climate variables with tree-ring formation. In this study, we applied the Vaganov-Shashkin model to simulate seasonal cell production and cell enlargement of Larix gmelinii Rupr. (Rupr.) growing in the permafrost zone of Central Siberia. We developed a procedure for calculating the radial cell diameter based on specific Gompertz function combined with the “instantaneous tracheidogram” approach to estimate effectively seasonal cell production and timing of cell enlargement under climate control. Simulated cell number and cell size matched well with direct xylogenesis observations. The developed procedure demonstrate strong relationships between seasonal simulated growth rate and measured tracheid radial size (the average correlation is 0.64, $p$ < 0.01). A highly significant correlation ($p$ < 0.01) between simulated and observed cell profiles was obtained for 71% of the growing seasons over the period 1950–2011. The strong exponential relationship (R² = 0.67) was obtained between the day of the year (DOY) when cambial cell transfers into enlargement zone and simulated time intervals of cell enlargement. Based on the strong exponential relationship it was possible to reproduce the basic pattern of the observed tracheidograms over 1950–2011 with a systematic overestimation of final cell sizes at the beginning of the growing season, which will be possible to eliminate by using more anatomical data (trees) and longer period. The proposed approach of simulating intra-annual cell dynamics (cell production) has a great potential for studying how climate affects tree-ring formation.     

An empirical approach to the analysis of forest stratification

Natural forest communities are made up of different overlapping elementary subpopulations consisting of individuals of different species and ages. To stratify the individuals of a forest stand into elementary subpopulations by using tree height records, a graphical method was empirically proposed employing a derivative of Pearson's type VII distribution as a basis. Arranging all of the individual trees in a stand in the descending order of their height (x), and introducing the rank (N) of any individual in the ordered ranking ofx, finite difference (n) ofN and three values ofx, i.e.,x(N), x(N−n), andx(N+n) labelled byN, N−n, andN+n, respectively, the proposed method used the linear relation betweenx(N) and [x(N−n)+x(N+n)]. On thex(N) vs.[x(N−n)+x(N+n)] diagram, the relation between both the variables could be approximated by a few segmental linear relations and used for stratifying the individuals of a forest stand into subpopulations. The method was applied for analyzing the vertical stratification in forest stands of different forest formations in Japan, Southeast Asia, Africa, and South America. The conclusions on stratification which resulted from the proposed method corresponded well to the conclusions on stratification from profile diagrams.     

A preliminary analysis of the DNA and diet of the extinct Beothuk: a systematic approach to ancient human DNA

We have used a systematic protocol for extracting, quantitating, sexing and validating ancient human mitochondrial and nuclear DNA of one male and one female Beothuk, a Native American population from Newfoundland, which became extinct approximately 180 years ago. They carried mtDNA haplotypes, which fall within haplogroups X and C, consistent with Northeastern Native populations today. In addition we have sexed the male using a novel-sexing assay and confirmed the authenticity of his Y chromosome with the presence of the Native American specific Y-QM3 single nucleotide polymorphism (SNP). This is the first ancient nuclear SNP typed from a Native population in the Americas. In addition, using the same teeth we conducted a stable isotopes analysis of collagen and dentine to show that both individuals relied on marine sources (fresh and salt water fish, seals) with no hierarchy seen between them, and that their water sources were pooled or stored water. Both mtDNA sequence data and Y SNP data hint at possible gene flow or a common ancestral population for both the Beothuk and the current day Mikmaq, but more importantly the data do not lend credence to the proposed idea that the Beothuk (specifically, Nonosabasut) were of admixed (European-Native American) descent. We also analyzed patterns of DNA damage in the clones of authentic mtDNA sequences; there is no tendency for DNA damage to occur preferentially at previously defined mutational hotspots, suggesting that such mutational hotspots are not hypervariable because they are more prone to damage.     

EMMLi: A maximum likelihood approach to the analysis of modularity

Identification of phenotypic modules, semiautonomous sets of highly correlated traits, can be accomplished through exploratory (e.g., cluster analysis) or confirmatory approaches (e.g., RV coefficient analysis). Although statistically more robust, confirmatory approaches are generally unable to compare across different model structures. For example, RV coefficient analysis finds support for both two‐ and six‐module models for the therian mammalian skull. Here, we present a maximum likelihood approach that takes into account model parameterization. We compare model log‐likelihoods of trait correlation matrices using the finite‐sample corrected Akaike Information Criterion, allowing for comparison of hypotheses across different model structures. Simulations varying model complexity and within‐ and between‐module contrast demonstrate that this method correctly identifies model structure and parameters across a wide range of conditions. We further analyzed a dataset of 3‐D data, consisting of 61 landmarks from 181 macaque (Macaca fuscata) skulls, distributed among five age categories, testing 31 models, including no modularity among the landmarks and various partitions of two, three, six, and eight modules. Our results clearly support a complex six‐module model, with separate within‐ and intermodule correlations. Furthermore, this model was selected for all five age categories, demonstrating that this complex pattern of integration in the macaque skull appears early and is highly conserved throughout postnatal ontogeny. Subsampling analyses demonstrate that this method is robust to relatively low sample sizes, as is commonly encountered in rare or extinct taxa. This new approach allows for the direct comparison of models with different parameterizations, providing an important tool for the analysis of modularity across diverse systems.     

The roof is on fire! A dendrochronological reconstruction of the restoration of the Basilica of Our Lady in Tongeren (Belgium)

A dendrochronological study of the roofs of the Basilica of Our Lady in Tongeren (Belgium) showed that the current roofs date to the period right after the deliberate burning of the church in 1677. High resolution dendrochronological dates – based on wood-anatomical observations of the outermost tree-ring – combined with detailed archival sources, allow us to trace the progress of reconstruction of the various roofs throughout the years following the fire at an intra-annual resolution. This provides an exceptionally detailed timeline of both the temporal and spatial progress of the reconstruction of the roofs of the Basilica of Our Lady. As such, this case-study can serve as a very detailed reference for the study of post-medieval roofs and the workflow during their construction.     

五种大戟属植物nrDNA的ITS序列分析及其叶的比较解剖学研究

运用PCR直接测序法和石蜡切片法,测定5种安徽产大戟属植物的nrDNA的内转录间隔区(ITS)序列包括5.8SrDNA及其叶片的解剖。结果表明(1)5种大戟属植物的ITS的长度范围为641~662bp,运用Maga软件构件ITS树把5种植物归为两大支大戟、月腺大戟与乳浆大戟聚为一支;地锦和斑地锦聚为另一支。结果表明地锦和斑地锦亲缘关系较近,而大戟、月腺大戟和乳浆大戟亲缘关系较近。(2)5种大戟属植物的叶片结构中,除地锦和斑地锦外,其余3种有明显的栅栏组织和海绵组织之分,但二者在叶肉中的厚度比在种间有一定的差别;而地锦和斑地锦叶的栅栏组织与海绵组织分化不明显,且皆有乳汁管,其他3种植物叶片内未见此结构,叶片结构分析表明大戟、月腺大戟和乳浆大戟叶片结构相近,而地锦草和斑地锦叶片结构也相近。(3)分析表明nrDNA的ITS序列及叶的比较解剖特征具有明显的相关性。以上研究结果为大戟属植物的分类和药用开发提供了实验证据。