Spatial and age-dependent tree-ring growth responses of Larix gmelinii to climate in northeastern China

Tree-ring width chronologies from 276 Larix gmelinii cores taken in northeastern China were used to analyze spatial and age-dependent growth–climate response relationships. Tree radial growth from five localities showed similar patterns, while exhibiting different tree-ring growth responses to local climate. The rotated principal component analysis (RPCA) indicated that tree age, growing season moisture conditions, and ambient air temperature variations resulted from location differences (e.g., longitude, latitude, and altitude), which could explain the non-stationary spatial climate–growth relations observed. The study tested the fundamental assumption that the climate–growth of L. gmelinii was age independent after the removal of size trends and disturbance signals. The age-related climate–growth relationship might potentially improve the veracity of past climate reconstructions. Bootstrapped correlation function analyses suggested that the response of L. gmelinii radial growth to climate differed between trees ≥150 years old and <150 years old. Mean sensitivity and standard deviation for trees increased with age in the <150 years old tree class; whereas trees ≥150 years old had no significant relationship with age. These results showed that the assumption of age-independent climate–growth relationship is invalid at these sites. Physiological processes and/or hydraulic constraints dependent on tree age, together with detrending techniques could be the possible causal factors of clear age-dependent responses. These results suggested the importance of incorporating trees of all ages into the chronology to recover a detailed climatic signal in a reconstruction of L. gmelinii for this region.     

Age-dependent tree-ring growth responses to climate in Qilian juniper (Sabina przewalskii Kom.)

Qilian juniper (Sabina przewalskii kom.) is one of the dominant tree species on Qinghai-Tibetan Plateau and has been used in dendroclimatological studies. Here we designed a test to examine whether or not the climate–growth responses in tree rings of Qilian Juniper vary with the change in tree's age. A total number of 135 increment cores were sampled from Qilian Juniper trees at five sites, in which 112 cores were selected and grouped into five 100-year age classes for analysis of age-dependent climate–growth relationships. Chronology statistics, response functions and ANOVA F-test were used to test the consistency of five age-class mean chronologies (AGCs). The results showed that mean sensitivity (MS) and standard deviation (SD) did not change significantly with age. Response function analysis indicated that (a) climate accounts for a high amount of variance in tree-ring widths; (b) tree-ring growth has significant positive correlation with mean monthly air temperature of previous October and November, and with total monthly precipitation of current January and June, while has significant negative correlation with mean monthly air temperature of current May; and (c) AGC-2, AGC-3 and AGC-4 have stronger response to climate change than AGC-1 and AGC-5. The ANOVA F-test showed that generally there are significant differences between the first age class and other four age classes, but among the four classes in which trees are older than 200 years, the differences are usually insignificant. Overall the long-lived Qilian Juniper is still an ideal tree species for dendroclimatic reconstruction.     

Correlation between Gliotoxin Production and Virulence of Aspergillus fumigatus in Galleria mellonella

Thanatephorus cucumeris is a ubiquitous fungus responsible for many types of plant diseases worldwide. All isolates from infected Hevea brasiliensis trees secreted pectolytic enzymes; polygalacturonase (PG), pectin lyase (PL) and cellulolytic enzymes; beta-glucosidase and cellobiase in culture. The extracts of the rubber tree leaf tissues, inoculated with T. cucumeris did not show any PG activity. However, PL activity was detected in tissue with the establishment of the infection. The levels of beta-glucosidase, an inherent enzyme in Hevea spp. increased rapidly following infection. However, cellobiase was detected only with the initiation of infection. Molecular weights of PG in all isolates were similar and in the range of 53,000 to 58,000. PL also followed the same pattern showing a molecular weight around 39,000.     

Purification and characterization of two isozymes of polygalacturonase from Botrytis cinerea. Effect of calcium ions on polygalacturonase activity

The phytopathogenic fungus Botrytis cinerea produces a set of polygalacturonases (PGs) which are involved in the enzymatic degradation of pectin during plant tissue infection. Two polygalacturonases secreted by B. cinerea in seven-day-old liquid culture were purified to apparent homogeneity by chromatography. PG I was an exopolygalacturonase of molecular weight 65 kDa and pI 8.0 and PG II was an endopolygalacturonase of 52 kDa and pI 7.8. Enzymatic activity of PG I and PG II was partially inhibited by 1 mM CaCl₂, probably by calcium chelation of polygalacturonic acid, the substrate of the enzyme.     

Male and Female Activity Patterns in Microcebus murinus During the Dry Season at Kirindy Forest, Western Madagascar

I investigated sexual differences in activity levels, and their adaptive significance in gray mouse lemurs, Microcebus murinus, during the dry season. I studied them between April and September, 1999, at Kirindy Forest, a dry deciduous forest in western Madagascar. Six males and 6 females were radiocollared and followed using focal individual sampling. The males remained active throughout the dry season, whereas the females hibernated for up to 3 mo. Males exhibited a gradual decrease in activity from May to June and some of them remained inactive for periods of a few days to 1 wk. Males were more active in July. Though there were changes in the duration of time spent sleeping, there was no significant change in male body mass throughout the period. Males appear to prefer to rest in nests that are close to or coincident with those preferred by females, and multiple males may simultaneously share nests with multiple hibernating females. Males occupying nests that are close to or coincident with nests preferred by females tend to switch nests less frequently than others do. Researchers have suggested that high levels of activity prepare males for the mating season, which occurs at the end of the dry season. Presumably, by maintaining an active lifestyle throughout the dry season, certain males may position themselves to monopolize the best tree holes during the mating season. The reproductive advantages gained by this behavior may overshadow the costs of remaining active during the dry season when resources are scarce, temperatures are low, and predator pressure is high. Measurements on captured males confirm a weak (nonsignificant) positive correlation between body mass and activity levels. There is no indication that the heavier, more active males were better able to monopolize the best tree holes.     

Regulation of synthesis of nitrite reductase in pea leaves: in-vivo and in-vitro studies

The cellular location of three peroxidase isoenzymes (PRX) in mature leaf tissue of Petunia and their affinity for Concanavalin A-Sepharose were investigated. The isoenzymes PRXa, PRXb and PRXc were identified by their positions in starch-gel zymograms. The fast-moving anodic and cathodic peroxidase bands, the isoenzymes PRXa and PRXc respectively, were the most active peroxidases in extracellular extracts. The molecular forms of PRXa showed a tissue-specific distribution between midrib and remaining leaf tissue. An intermediate-moving anodic peroxidase band, the isoenzyme PRXb, was the most active peroxidase released after extraction of isolated mesophyll protoplasts. Small amounts of the peroxidase isoenzymes were present in cell-wall-bound fractions. Incubation of a crude protein fraction with Concanavalin A-Sepharose showed that the isoenzyme PRXb bound more firmly to Concanavalin A-Sepharose than the isoenzymes PRXa and PRXc, of which only one molecular form bound partly. The results are discussed with respect to a possible function of one of the peroxidase isoenzymes, and a possible role of oligosaccharide chains in determining the cellular location of plant peroxidases is suggested.Abbreviations Con A Concanavalin A - PRX peroxidase (isoenzyme)     

Variations in the population structure and ecology of Matteuccia struthiopteris

The aim of the study was to investigate variation in the structure of a population of Matteuccia struthiopteris (L.) Tod. in N–E Poland depending on its location in a river valley and light availability under tree canopies. Ordination statistical methods and structural equation modeling were used to identify ecological relationships in the population. The analysis showed that optimum growth conditions of M. struthiopteris depend on the canopy cover. As the shading increased, the size of trophophyllous leaves and the production of sporophylls decreased. The relationship between the canopy cover and the optimum growth conditions was best expressed by rootstock diameter and, to a lesser extent, by the number and height of trophophylls. An indirect relationship between the number and height of sporophylls and the canopy cover, by rootstock size and trophophyll quantity, was also identified. This relationship may be caused by the predominance of vegetative forms of reproduction. Based on the correlations between the landscape position vs. rootstock diameter and trophophyll height, a difference in the habitat colonization potential of M. struthiopteris could be detected. Three spatial clusters of M. struthiopteris in the river valley were identified: river bank, river terrace, river valley slopes. These relationships and the distribution of other herbaceous plant species occurring at the plots indicate that the distribution pattern of M. struthiopteris in the study area is determined by diverse environmental conditions, however mainly by light conditions.     

Changes in leaf morphology and anatomy with tree age and height in the broadleaved evergreen species, Eucalyptus regnans F. Muell

Relatively little is known about changes in leaf attributes over the lifespan of woody plants. Knowledge of such changes may be useful in interpreting physiological changes with age. This study investigated changes in leaf morphology and anatomy with tree age and height in the broadleaved evergreen species, Eucalyptus regnans. Fully expanded leaves were sampled from the upper canopy of tree ages ranging from 6 to 240 years, and tree heights ranging from about 10–80 m. There were significant changes in leaf form with increasing tree age and height. Leaf size and specific leaf area (SLA; leaf area/leaf mass) decreased, leaf thickness increased, and leaves became narrower relative to their length, with increasing tree age and height. Cuticle thickness and leaf waxiness, including wax occlusion of the stomatal antechamber, increased with increasing age and height. By comparison, there were no clear trends in stomatal frequency or stomatal length with tree age, although there were curvilinear relationships between an index of total stomatal pore area per leaf lamina and both tree age and tree height. The results support the hypothesis that leaves of E. regnans become more xeromorphic with tree age and height. The results are discussed in relation to their significance for changes in water relations in the canopy with age.     

Interaction between the root-lesion nematode Pratylenchus vulnus and the mycorrhizal association of Glomus intraradices and Santa Lucia 64 cherry rootstock

The effects of the interaction between Pratylenchus vulnus and the endomycorrhizal fungus Glomus intraradices on growth and nutrition of Santa Lucia 64 cherry rootstock was studied under microplot conditions during one growing season. Fresh top weight, and stem diameter of mycorrhizal plants and high P treatments with and without P. vulnus were significantly higher than those of non-mycorrhizal plants. The lowest shoot length and fresh root weights were recorded in nematode inoculated plants in low P soil. Mycorrhizal infection did not affect the number of nematodes per gram of root in plants infected with P. vulnus. In the presence of the nematode, internal spore production by G. intraradices was significantly reduced. No nutrient deficiencies were detected through foliar analysis, although low levels of Ca, Mn and Fe were detected in nematode treatments. Mycorrhizal plants achieved the highest values for N, P, S, Fe, and Zn, whereas high P treatments increased absorption of Ca and Mn. Early mycorrhizal infection of Santa Lucia 64 cherry rootstock by G. intraradices confers increased growth capacity in the presence of P. vulnus.     

Phytophthora collar rot of apple: influence of the rootstock on scion variety resistance

In field experiments with young trees great differences were found in the resistance to Phytophthora cactorum of Cox's Orange Pippin apple scions grafted on different clonal rootstocks. The rootstock effect on scion resistance was inversely related to the effect on tree vigour: the rootstocks inducing high resistance were dwarfing (M. 9) or semi-dwarfing (M. 7, M. 26, MM. 106), and those inducing low resistance were vigorous or very vigorous (M. 25, MM. 104, MM. 109). Mean lesion lengths in Cox on MM. 104 were five to eight times greater than those in Cox on M. 9. The rootstock influence on scion resistance was associated primarily with effects on the rate of lesion extension: during the early stages only of host colonization there appeared to be threshold extension rates below which host resistance factors effectively suppressed a large proportion of infections. The influence of the root-stock on scion resistance was apparently unrelated to inherent rootstock resistance. On all rootstocks Cox showed diminished resistance to infection during the period from the swelling of buds to the early stages of shoot growth. Although most susceptible during the ‘mouse-ear’ and ‘pink bud’ stages of development, suscpetibility was not associated with flowering per se. Rootstock type did not affect the resistance of Cox scions to P. syringae, for which the period of susceptibility to infection occurred in the dormant season.     

Investigating the effects of age‐related spatial structuring on the transmission of a tick‐borne virus in a colonially breeding host

Higher pathogen and parasite transmission is considered a universal cost of colonial breeding due to the physical proximity of colony members. However, this has rarely been tested in natural colonies, which are structured entities, whose members interact with a subset of individuals and differ in their infection histories. We use a population of common guillemots, Uria aalge, infected by a tick‐borne virus, Great Island virus, to explore how age‐related spatial structuring can influence the infection costs borne by different members of a breeding colony. Previous work has shown that the per‐susceptible risk of infection (force of infection) is different for prebreeding (immature) and breeding (adult) guillemots which occupy different areas of the colony. We developed a mathematical model which showed that this difference in infection risk can only be maintained if mixing between these age groups is low. To estimate mixing between age groups, we recorded the movements of 63 individually recognizable, prebreeding guillemots in four different parts of a major colony in the North Sea during the breeding season. Prebreeding guillemots infrequently entered breeding areas (in only 26% of watches), though with marked differences in frequency of entry among individuals and more entries toward the end of the breeding season. Once entered, the proportion of time spent in breeding areas by prebreeding guillemots also varied between different parts of the colony. Our data and model predictions indicate low levels of age‐group mixing, limiting exposure of breeding guillemots to infection. However, they also suggest that prebreeding guillemots have the potential to play an important role in driving infection dynamics. This highlights the sensitivity of breeding colonies to changes in the behavior of their members—a subject of particular importance in the context of global environmental change.     

Variation of adult Great Tit Parus major body condition and blood parameters in relation to sex, age, year and season

In recent years, an increasing number of studies have focused on the ranges of variation of health related biochemical and haematological parameters in wildlife, but information is still scarce for enzymatic activities which can be extremely important in detecting potential responses to environmental change. In a Great Tit (Parus major) population, we describe the variation in relation to age, sex, season and year of: (1) morphological: body condition index, fat and muscle, (2) haematological: hematocrit, haemoglobin, white blood cell count and heterophil/lymphocyte ratio, and (3) biochemical parameters: plasma protein and activities of plasma cholinesterases (acetylcholinesterase and butyrylcholinesterase) activity and red blood cell glutathione peroxidase. Sex had significant effects on all morphological parameters except fat. Age significantly affected cholinesterase activities, H/L ratio and haemoglobin, and there was a significant interaction between sex and age affecting hematocrit. There were significant interactions of year and season affecting almost all parameters studied—body condition index, fat, protein, acetyl and butyrylcholinesterase activities, glutathione peroxidase activity and haemoglobin. This study indicates that these parameters are largely influenced by year and seasonal effects, besides the individual’s intrinsic variation. Therefore, when evaluating experimental or environmental change effects, appropriate controls should be used.     

Seasonal Effects on Sleeping Site Ecology in a Nocturnal Pair-Living Lemur (Avahi occidentalis)

Seasonal changes may have a strong effect on the safety of sleeping sites in arboreal primates. For example, changes in vegetation thickness may impact predation risk and energy expenditure related to thermoregulation. We investigated how seasonality influenced sleeping site characteristics and usage pattern in an arboreal primate living in a highly seasonal environment. The western woolly lemur (Avahi occidentalis) lives in the dry deciduous forest of northwestern Madagascar, where leaf coverage greatly varies across the year. We examined the hypothesis that these lemurs change their sleeping site behavior dependent on season. We collected data on sleeping site height and location, and characterized usage patterns in six radiotagged pairs between May and December 2008. During the late dry season, pairs preferentially slept in the middle part of a tree. In contrast, there was no height preference during the early rainy season. The lemurs used more sleeping sites during the early rainy than during the late dry season and stayed more days at the same sleeping tree in the late dry season. Our findings support the hypothesis that season affects sleeping site selection in an arboreal primate species living in a highly seasonal environment. During the late dry season, western woolly lemurs are particularly conspicuous to hunters and we therefore suggest a better monitoring of the forest in this season to guarantee their future survival.     

Fine root architecture, morphology, and biomass of different branch orders of two Chinese temperate tree species

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China—Larix gmelinii Rupr and Fraxinus mandshurica Rupr —by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and.10–20 cm). Branching ratio (R _b) differed with the level of branching: R _b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R _b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.     

Roots,soil water and crop yield: tree crop interactions in a semi-arid agroforestry system in Kenya

Variations in soil water, crop yield and fine roots of 3–4 year-old Grevillea robusta Cunn. and Gliricidia sepium (Jacq.) Walp. growing in association with maize (Zea mays L.) were examined in semiarid Kenya during the long rains of 1996 and 1997. Even although tree roots penetrated more deeply than maize roots, maximum root length densities for both tree species and maize occurred in the top 200 mm of the soil profile where soil moisture was frequently recharged by rains. Populations of roots in plots containing trees were dominated by tree roots at the beginning of the growing season but because tree roots died and maize root length increased during the cropping season, amounts of tree and maize roots were similar at the end of the season. Thus, there was evidence of temporal separation of root activity between species, but there was no spatial separation of the rooting zones of the trees and crops within that part of the soil profile occupied by crop roots. Tree root length density declined with increasing distances from rows of trees and with depth in the soil profile. Although Grevillea trees were largest, plots containing G. sepium trees always contained more tree roots than plots containing G. robusta trees and Gliricidia was more competitive with maize than Grevillea. Overall, Gliricidia reduced crop yield by about 50% and Grevillea by about 40% relative to crop yield in control plots lacking trees and reductions of crop yield were greatest close to trees. There was less soil moisture in plots containing trees than in control plots. Such difference between control plots and plots containing trees were maximal at the end of the dry season and there was always less soil moisture close to trees than elsewhere in the plots. Plots containing Gliricidia trees contained less soil water than plots containing Grevillea trees.     

Effects of extracts of Alnus glutinosa seeds on growth of Frankia strain ArI3 under static and fermentor culture conditions

The amount of root mortality caused by root pathogens such as Phytophthora nicotianae (syn. Phytophthora parasitica) has typically been inferred from the net change in root length density in sequential soil cores. Because such measurements give information only on net changes in root populations, the actual rate of root turnover is often underestimated. We used minirhizotrons to track the fate of a large number of individual fine roots of mature field-grown citrus trees over a 6-month period. This method enabled us to examine the effect of P. nicotianae population levels on fine-root mortality. Seasonal and genotypic variation in patterns of citrus fine root mortality were associated with variation in population levels of P. nicotianae. Fine root lifespans were shorter when populations of P. nicotianae were high. Fine roots of the Phytophthora-susceptible rootstock, rough lemon (Citrus jamibhiri), had shorter median lifespans and supported larger populations of P. nicotianae than the fine roots of the more tolerant rootstock, Volkamer lemon (Citrus volkameriana). Rates of root mortality were either relatively constant for roots of all ages, or increased with age; the latter pattern was most pronounced for Volkamer lemon roots. Differences in the age-dependence of root mortality may, therefore, play a role in genotypic differences in tolerance of Phytophthora root rot by these two rootstocks. H Lambers Section editor     

Age-dependent responses of tree-ring growth and intra-annual density fluctuations of Pinus pinaster to Mediterranean climate

Dendrochronology generally assumes that climate–growth relationships are age independent once the biological growth trend has been removed. However, tree physiology, namely, photosynthetic capacity and hydraulic conductivity changes with age. We tested whether the radial-growth response to climate and the intra-annual density fluctuations (IADFs) of Pinus pinaster Ait. varied with age. Trees were sampled in Pinhal de Leiria (Portugal), and were divided in two age classes: young (<65 years old) and old (>115 years old). Earlywood and tree-ring width of young P. pinaster trees were more sensitive to climate influence while the response of latewood width to climate was stronger in old trees. Young trees start the growing season earlier, thus a time window delay occurs between young and old trees during which wood cells of young trees integrate environmental signals. Young trees usually have a longer growing season and respond faster to climate conditions, thus young P. pinaster trees presented a higher frequency of IADFs compared with old trees. Most of the IADFs were located in latewood and were positively correlated to autumn precipitation. The radial-growth response of P. pinaster to climate and the IADFs frequency were age dependent. The use of trees with different age to create a tree-ring chronology for climate studies can increase the resolution of climatic signals. Age-dependent responses to climate can also give important clues to predict how young and old trees react to climate change.     

Fine-scale distribution of treeline trees and the nurse plant facilitation on the eastern Tibetan Plateau

Above-average climate warming occurred during the 20th century in high altitude regions, and alpine treelines are believed to be an early indicator to respond to these warming-related changes. However, empirical investigations on treeline dynamics showed diverse results. The main objectives of this study are: (1) to investigate if treeline position shifted and if tree recruitment changed along with climate warming, and (2) to test if adult trees have “nursing effect” on tree establishment at treelines. We investigated two Balfour spruce (Picea balfouriana Rehd. et Wils.) treelines in Chang Niang (CNT) and Dang Dui (DDT), Dingqing county, Changdu prefecture, eastern Tibet. At each treeline site, three replicate plots with a size 30 m × 50 m were established. The coordinates of each tree within the plots were recorded and the age of each tree was identified by dendrochronological method. The changes in treeline position and tree recruitment were examined from spatially fine-scale distribution of trees and their age structure. The spatial patterns of individual trees were analyzed to infer the neighborhood effects. Results indicate that plots CNT2, CNT3, DDT1 and DDT2 showed stable treeline position during the last century, whereas plots CNT1 and DDT3 showed treeline advancing movement. Tree recruitments in all the six plots were enhanced during the 20th century, with two peaks occurring in the 1890–1910s and the 1950–1990s. Seedlings and saplings showed a general clustered distribution in all the six plots. The diverse pattern of treeline movement and episodic regeneration suggest that the treeline activity is not merely a result of climate change. “Nursing effects” from adult trees may play an important role in shaping the treeline activities on the eastern Tibetan Plateau. Our findings reveal diverse patterns in treeline dynamics at a local scale and highlight the importance of incorporating biotic interactions into species distribution modeling approaches.     

Modelling variability of wood density in beech as affected by ring age,radial growth and climate

Although it has been recognized as a key parameter of wood quality and a good source of information on growth, annual wood density has been little studied within diffuse-porous trees such as beech ( Fagus sylvatica Liebl.). In this paper we examine the variability encountered in beech ring density series and analyze the influences of ring age, ring width, climate and between-tree variability on density. Thirty ring sequences were sampled from 55-year- old dominant beech trees growing within the same stand; ring density and width were measured using radiography. Ring density proved to be less variable through time than ring width. The relationship between these two variables was less than observed in ring-porous trees and it showed great variation between trees. The sensitivity of ring width and density to climate was also different; width was strongly linked to soil water deficit whereas density was correlated to temperature and August rainfall. Unlike ring width, wood density showed sensitivity towards climatic characteristics of the late growing season. A large part of annual density variability remains unexplained, even using advanced modelled water balance variables. We hypothesize that a significant part of the tree ring is under internal control. We also demonstrated great inter-tree variability (the tree effect) in ring density, which has an influence on density but not on trees response to climate.