Effect of ecosystem warming on boreal black spruce bud burst and shoot growth

The boreal forest is predicted to experience the greatest warming of any forest biome during the next 50–100 years, but the effects of warming on vegetation phenology are not well known. The objectives of this study were to (1) examine the effects of whole ecosystem warming on bud burst and annual shoot growth of black spruce trees in northern Manitoba, Canada and (2) correlate bud burst to cumulative degree-days (CDD). The experimental design was a complete randomized block design that consisted of four replicated blocks. Each replicate block contained four treatments: soil warming only (heated outside, HO), soil and air warming (heated inside, HI), control outside (no chamber, no heating, CO), and inside a chamber maintained at ambient conditions (no soil or air warming, control inside, CI). Bud burst was measured during the first and second years of the experiment, starting in 2004, and annual shoot growth was measured for the first 3 years (2004–2006) of the study. On average, shoot bud burst occurred 11 and 9 days earlier in 2004 and 2005, respectively, for HI than for other treatments. However, mean CDD required for bud burst for HI was within the standard deviation of CO for both years. In year 1 of the treatments, shoot bud burst occurred earlier for HI than other treatments (CI, CO, HO), but final shoot length of HI trees was less than in CO trees. In the second year of warming, final shoot length was not different for HI than CO. By the third year of warming final shoot length was significantly greater for HI than all other treatments. Empirical results from this study suggest that soil and air warming causes an earlier bud burst for all years of observation and greater shoot lengths by the third season of warming. A longer growing season and greater annual shoot growth should increase carbon uptake by boreal black spruce trees in a warmer climate.     

Comparing growth phenology of co-occurring deciduous and evergreen conifers exposed to drought

Plant phenological events are influenced by climate factors such as temperature and rainfall. To evaluate phenological responses to water availability in a Spring Heath-Pine wood (Erico-Pinetum typicum), the focus of this study was to determine intra-annual dynamics of apical and lateral growth of co-occurring early successional Larix decidua and Pinus sylvestris and late successional Picea abies exposed to drought. The effect of reduced plant water availability on growth phenology was investigated by conducting a rainfall exclusion experiment. Timing of key phenological dates (onset, maximum rate, end, duration) of growth processes were compared among species at the rain-sheltered and control plot during 2011 and 2012. Shoot and needle elongation were monitored on lateral branches in the canopy at c. 16 m height and radial growth was recorded by automatic dendrometers at c. 1.3 m height of >120 yr old trees. Different sequences in aboveground growth phenology were detected among the three species under the same growing conditions. While onset of radial growth in April through early May was considerably preceded by onset of needle growth in Larix decidua (5–6 weeks) and shoot growth in Pinus sylvestris (c. 3 weeks), it occurred quite simultaneously with onset of shoot growth in Picea abies. Low water availability had a minor impact on onset of aboveground growth, which is related to utilization of stored water, but caused premature cessation of aboveground growth. At the control plot mean growing season length was 130 days in Pinus sylvestris, 95 days in Larix decidua and 73 days in Picea abies, supporting the hypothesis that early successional species are resource expenders, while late successional species are more efficient in utilizing resources and develop safer life strategies. High synchronicity found in culmination of radial growth in late spring (mid-May through early June) prior to occurrence of more favourable environmental conditions in summer might indicate sink competition for carbohydrates to belowground organs. This is supported by completion of apical growth in mid June in all species, except for needle growth of Pinus sylvestris which lasted until early August. Phenological observations of conifers exposed to drought revealed that tree water status early during the growing season determines total annual aboveground growth and, besides temperature, species-specific endogenous and/or environmental factors (most likely photoperiod and/or different threshold temperatures) are involved in controlling apical and lateral growth resumption after winter dormancy.     

A model including photoperiod in degree days for estimating Hevea bud growth

 The aim of this study was to verify the applicability of a heat-sum model and an alternative model that included photoperiod to describe Hevea bud growth. Considering 19° C as the base temperature, approx. 172 degree days were required for bud growth up to leaf blade expansion. The applicability of the heat-sum model for Hevea bud growth was confirmed. The inclusion of day length as an exponential term in accumulated degree days resulted in the model fitting much better for the spring than for the winter conditions. Received: 21 May 1995 / Revised: 28 October 1996 / Accepted: 27 January 1997     

Regional trends for bud burst and flowering of woody plants in Norway as related to climate change

Data series for bud burst, beginning of flowering and petal fall for 20 species of deciduous trees and conifers at four sites in different regions of southern Norway have been analysed and related to temperature series. On average, the spring phenophases occurred 7 days earlier during the period 1971-2005. The most significant linear trends were observed for the earliest phases. The trends in this period were compared with trends in other periods, the longest one starting in 1927. Those starting in cold decades and ending in 2005 were in most instances statistically significant, whereas hardly any significant trend appeared for series starting in warm decades. This fact showed that the results of trend studies are very sensitive to the choice of starting year. There were significant decadal variations in 40% of the series. The dates of occurrence of the phenophases, varying from the first days of May to the first days of June, correlated with seasonal temperature series, in most cases strongest to mean temperatures for the seasons March-May and April-May. The North Atlantic Oscillation Index (NAOI) for January and February appeared to have some predictive power for the date of occurrence of the recorded phases. The basis for this may be that the oscillations described by the index are of importance for the fulfilment of physiological chilling requirements needed to break bud dormancy. The same genotypes of the trees were grown in region West Norway and in Central Norwegian region; during the period 1965-2005 the trends towards earlier bud burst were more pronounced and steeper at the western site.     

Increases in cytokinin levels in Scots pine in relation to chilling and bud burst

Levels of isopentenyladenosine and zeatin riboside were monitored in buds and needles of Scots pine ( Pinus sylvestris L.) seedlings grown under controlled climatic conditions and in field-grown trees. Extracts were purified by immunoaffinity chromatography and high-performance liquid chromatography. Cytokinin levels were quantified with an enzyme-linked immunosorbent assay. The cytokinin content was low in buds and needles of dormant seedlings but increased during dormancy release, reaching peak values in buds just before resumption of shoot growth. Samples collected in the field also showed a marked increase in the levels of cytokinins just prior to bud burst. Further, the biological activity of applied cytokinins in activating the dormant buds of Scots pine is shown. The results indicate a probable role of cytokinins in seedlings during dormancy release.     

Spatio-temporal relief from hypoxia and production of reactive oxygen species during bud burst in grapevine (Vitis vinifera)

Background and Aims Plants regulate cellular oxygen partial pressures (pO₂), together with reduction/oxidation (redox) state in order to manage rapid developmental transitions such as bud burst after a period of quiescence. However, our understanding of pO₂ regulation in complex meristematic organs such as buds is incomplete and, in particular, lacks spatial resolution.Methods The gradients in pO₂ from the outer scales to the primary meristem complex were measured in grapevine (Vitis vinifera) buds, together with respiratory CO₂ production rates and the accumulation of superoxide and hydrogen peroxide, from ecodormancy through the first 72 h preceding bud burst, triggered by the transition from low to ambient temperatures.Key Results Steep internal pO₂ gradients were measured in dormant buds with values as low as 2·5 kPa found in the core of the bud prior to bud burst. Respiratory CO₂ production rates increased soon after the transition from low to ambient temperatures and the bud tissues gradually became oxygenated in a patterned process. Within 3 h of the transition to ambient temperatures, superoxide accumulation was observed in the cambial meristem, co-localizing with lignified cellulose associated with pro-vascular tissues. Thereafter, superoxide accumulated in other areas subtending the apical meristem complex, in the absence of significant hydrogen peroxide accumulation, except in the cambial meristem. By 72 h, the internal pO₂ gradient showed a biphasic profile, where the minimum pO₂ was external to the core of the bud complex.Conclusions Spatial and temporal control of the tissue oxygen environment occurs within quiescent buds, and the transition from quiescence to bud burst is accompanied by a regulated relaxation of the hypoxic state and accumulation of reactive oxygen species within the developing cambium and vascular tissues of the heterotrophic grapevine buds.     

植物激素对空心泡叶片不定芽形成的影响

空心泡是蔷薇科悬钩子属植物,属小果类果树,其聚合果肉质多浆,芳香,果味酸甜。空心泡目前尚处于野生状态,是一种很有利用前景的植物。有关悬钩子属植物组织培养已有许多报道,所有这些工作都     

Cytokinin effect on branching and bud formation in Funaria

Cytokinins have two different effects on protonemata of Funaria hygrometrica. They induce branching of unbranched caulonemata and bud formation. Branching occurs after treatment with pico-molar concentrations of cytokinins whereas bud formation requires micro-molar concentrations. Both processes are therefore independently stimulated by cytokinins.Abbreviation BA N⁶-benzyladenine     

Crown architecture of grafted Stone pine (Pinus pinea L.): shoot growth and bud differentiation

The singular umbrella-like crown shape of Stone pine can be interpreted as a consequence of primary shoot-growth patterns and posterior axis differentiation due to differential secondary growth and down-bending of branches. This paper centres on the first aspect, analysing the growth, branching and flowering behaviour of about 5,000 individual shoots on 27 grafted Stone pines. The data measurement on standing trees allowed to study correlations of topologic and geometric variables in the shoot and their ancestors. The only significant correlations were found with parameters of the mother shoot formed the previous year and with the number of cones born 3 years before by the respective ancestor. The fitted relationships between geometric and topologic shoot and branch variables are the first step of a structural model construction that can be completed with functional components like a radiation and a carbon allocation submodel, stressing the importance of the heavy Stone pine cones as carbon sinks, with a total annual allocation similar to stem wood. In conclusion, the Stone pine crown shape emerges as consequence of the lack of initial vigour differentiation between stem and main-branch apical meristems that favour the generalized sylleptic reiteration in the open-grown trees.     

Changing relationships between tree growth and climate in Northwest China

Recently, several studies have shown changing relationships between tree growth and climate factors, mostly in the circumpolar north. There, changing relationships with climate seem to be linked to emergent subpopulation behavior. Here, we test for these phenomena in Northwest China using three tree species (Pinus tabulaeformis, Picea crassifolia, and Sabina przewalskii) that had been collected from six sites at Qilian Mts. and Helan Mts. in Northwest China. We first checked for growth divergence of individual sites and then investigated the relationship between tree growth and climate factors using moving correlation functions (CF). Two species, Pinus and Sabina, from two sites clearly showed growth divergence, not only in the late twentieth century as reported in other studies, but also over nearly the whole record. In divergent sites, one chronology shows more stable relationships with climate factors (usually precipitation). In non-divergent sites, nearly all relationships either vary in strength or become non-significant at one point. While this might possibly be related to increased stress on some trees due to increasing temperature, the exact causes for this shift in sensitivity remain unclear. We would like to highlight the necessity for additional studies investigating possible non-stationary growth responses of trees with climate, especially at sites that are used for climate reconstruction as our sites in Northwest China.     

TWIG: a model to simulate the gravitropic response of a tree axis in the frame of elasticity and viscoelasticity, at intra-annual time scale

Trees are able to maintain or modify the orientation of their axes (trunks or branches) by tropic movements. For axes in which elongation is achieved but cambial growth active, the tropic movements are due to the production of a particular wood, called reaction wood which is prestressed within the growing tree. Several models have been developed to simulate the gravitropic response of axes in trees due to the formation of reaction wood, all within the frame of linear elasticity and considering the wood maturation as instantaneous. The effect viscoelasticity of wood has, to our knowledge, never been considered. The TWIG model presented in this paper aims at simulating the gravitropic movement of a tree axis at the intra-annual scale. In this work we studied both the effect of a non-instantaneous maturation process and of viscoelasticity. For this purpose, we considered the elastic case with maturation considered as an instantaneous process as the reference. The introduction of viscoelasticity in TWIG has been done by coupling TWIG to a model developed for bridges. Indeed from a purely mechanical point of view, bridges and trees are very similar: they are structures which are built in stages, they are made of several materials (composite structures), their materials are prestressed (wood is prestressed during the maturation process as a result of polymerisation of lignin and cellulose to form the secondary cell wall and concrete is prestressed during drying). Simulations gave evidence that the reorientation process of axes can be significantly influenced by the kinetics of maturation. Moreover the model has now to be tested with more experimental data of wood viscoelasticity but it appears that in the range of a relaxation time from 0 to 50 days, viscoelasticity has an important effect on the evolution of tree shape as well as on the values of prestresses.     

The ecological significance of phenology in four different tree species: effects of light and temperature on bud burst

The process of adaptation is the result of stabilising selection caused by two opposite forces: protection against an unfavourable season (survival adaptation), and effective use of growing resources (capacity adaptation). As plant species have evolved different life strategies based on different trade offs between survival and capacity adaptations, different phenological responses are also expected among species. The aim of this study was to compare budburst responses of two opportunistic species (Betula pubescens, and Salix x smithiana) with that of two long-lived, late successional species (Fagus sylvatica and Tilia cordata) and consider their ecological significance. Thus, we performed a series of experiments whereby temperature and photoperiod were manipulated during dormancy. T. cordata and F. sylvatica showed low rates of budburst, high chilling requirements and responsiveness to light intensity, while B. pubescens and S. x smithiana had high rates of budburst, low chilling requirements and were not affected by light intensity. In addition, budburst in B. pubescens and S. x smithiana was more responsive to high forcing temperatures than in T. cordata and F. sylvatica. These results suggest that the timing of growth onset in B. pubescens and S. x smithiana (opportunistic) is regulated through a less conservative mechanism than in T. cordata and F. sylvatica (long-lived, late successional), and that these species trade a higher risk of frost damage for the opportunity of vigorous growth at the beginning of spring, before canopy closure. This information should be considered when assessing the impacts of climate change on vegetation or developing phenological models.     

Phenology of nutritional differences between new and mature leaves and its effect on caterpillar growth

Abstract. 1. We determined the phenology of the shrub Spiraea latifolia Ait. Bork. (Rosaceae), which has indeterminate shoot growth, and the effects of phenological changes in leaf quality on growth rate of the early-spring feeding buckmoth caterpillars, Hemileuca lucina Hy. Edw. (Saturniidae).
2. Leaves, regardless of whether they were newly expanded or several weeks old, were tougher later in the growth season (mid-June) than similarly aged leaves collected earlier; correspondingly, water and nitrogen content for leaves of all ages declined through the larval period. By July, newly expanded leaves had no more nitrogen than mature leaves.
3. Relative growth rate of third instar larvae fed new leaves or a mixture of new and mature leaves in early June was higher than that of those fed mature leaves, and efficiency of conversion of digested food to biomass was higher for larvae fed new leaves than for those fed mature leaves or a mixture.
4. In another experiment, larvae were reared on new leaves through the fourth instar and then fed a diet of new, mature or a combination of new and mature leaves, a regimen that was similar to the phenologies of both plants and caterpillars in the field. There was no difference in time to pupation or pupal weights among these treatments.     

Growth responses of an African savanna tree, <Emphasis Type="Italic">Bauhinia thonningii</Emphasis> Schumacher,to defoliation,fire and climate

This study investigated the growth responses to defoliation, fire-exposure and climate factors of a widespread Africana savanna tree, Bauhinia thonningii Schumacher, at a site in central Zambia. Experimental trees (n = 47) were either exposed to fire (n = 12) in the first half of the dry season or protected from fire (n = 35). Some of the fire-protected trees (n = 12) were subjected to artificial defoliation in two consecutive years. Phenological responses (bud break, leaf flush and leaf production) to fire-exposure and defoliation were monitored on permanently marked sample shoots over a 2-year period. Radial tree growth (diameter at 1.3 m above ground) was measured annually over a 7-year period from 1998 to 2005. During the first two years, defoliation and fire-exposure advanced the onset of bud break and leaf flush but fire-exposed trees produced significantly less leaves than did trees protected from fire, probably because scorching caused more severe shoot die-back than is normal. Leaf production was also significantly affected by experimental treatments and their interaction with year. Although treatments had significant short-term effects on radial growth, previous-year growth significantly influenced current-year growth, thereby confirming the existence of autocorrelation in the time-series growth data of B. thonningii. The interaction between previous-year growth and climate factors explained a significant proportion (25–40%, P < 0.001) of the variance in annual tree growth. Variogram models predicted that a 2-year manual defoliation treatment would shorten the longer-term growth cycle while continuous fire-exposure extended the cycle by one year. The results are useful for the management of savanna trees.     

A mathematical model to describe the dynamic response of a spruce tree to the wind

 A mathematical, computer-based, dynamic sway model of a Sitka spruce (Picea sitchensis) tree was developed and tested against measurements of the movement of a tree within a forest. The model tree was divided into segments each with a stiffness, mass and damping parameter. Equations were formulated to describe the response of every segment which together form a system of coupled differential equations. These were solved with the aid of matrices and from the resulting modes, the transfer function of the tree was found and used to calculate the movement of the tree in the wind. Comparison of the modelled movement of a tree in response to the measured wind speed above a forest canopy gave good agreement with the measured movement of the top of the tree but less satisfactory agreement close to the base. The comparison also pointed to the complexity of tree response to the wind and inadequacies in the model. In particular, the branches need to be treated as coupled cantilevers attached to the stem rather than simply as masses lumped together. Received: 18 February 1997 / Accepted: 16 December 1997     

Climatic effect of atmospheric CO2 doubling on radial tree growth in south eastern France

The climatic effect of a doubling of atmospheric CO₂ on radial growth of trees was studied in ten populations of three species in south eastern France using an Atmospheric General Circulation Model (AGCM) predicting a 3°C increase of mean temperature and a light rise of precipitation. Results are based on empirical growth climate models, involving an Artificial Neural Network (ANN) technique. Only two of the studied populations, on the boundaries of their ecological area, are sensitive to the climatic variations. One is the larch ( Larix decidua Mill.) population located at 2300 m on elevation (near the timberline) which shows a radial growth increase. The other is the most southern French Scots pine ( Pinus sylvestris L.) population which reacts with a severe growth rate reduction.     

Canopy structures and its effect on shoot growth and flowering in subalpine forests

Changes in foliage density distribution with altitude and its effect on shoot growth and flowering were studied in forest section planes (profiles) of subalpine forests and scrubs (Krummholz) in Nepal and Japan.Patterns of foliage in forest canopy were evaluated by an analysis of variance. Foliage densities were very high at high altitude canopies, but the change in spatial patterns along altitude differs in both areas.The spatial pattern of new shoot production was similar to that of current foliage density and was affected by the amount of foliage above the sample probably through light condition. Flowering of tall trees occurred in the surface of the upper canopy, but a shrub species flowered even under tree canopies.