Stem Girdling Research Articles

Stem girdling affects the onset of autumn senescence in aspen in interaction with metabolic signals.

Autumn senescence in aspen (Populus tremula) is precisely timed every year to relocate nutrients from leaves to storage organs before winter. Here we demonstrate how stem girdling, which leads to the accumulation of photosynthates in the crown, influences senescence. Girdling resulted in an early onset of senescence, but the chlorophyll degradation was slower and nitrogen more efficiently resorbed than during normal autumn senescence. Girdled stems accumulated or retained anthocyanins potentially providing photoprotection in senescing leaves. Girdling of one stem in a clonal stand sharing the same root stock did not affect senescence in the others, showing that the stems were autonomous in this respect. One girdled stem with unusually high chlorophyll and nitrogen contents maintained low carbon‐to‐nitrogen (C/N) ratio and did not show early senescence or depleted chlorophyll level unlike the other girdled stems suggesting that the responses depended on the genotype or its carbon and nitrogen status. Metabolite analysis highlighted that the tricarboxylic acid (TCA) cycle, salicylic acid pathway, and redox homeostasis are involved in the regulation of girdling‐induced senescence. We propose that disrupted sink‐source relation and C/N status can provide cues through the TCA cycle and phytohormone signaling to override the phenological control of autumn senescence in the girdled stems.

Predicting tropical tree mortality with leaf spectroscopy

AbstractDo tropical trees close to death have a distinct change to their leaf spectral signature? Tree mortality rates have been increasing in tropical forests, reducing the global carbon sink. Upcoming hyperspectral satellites could be used to predict regions close to experiencing extensive tree mortality during periods of stress, such as drought. Here we show, for a tropical rainforest in Borneo, how imminent tropical tree mortality impacts leaf physiological traits and reflectance. We measured leaf reflectance (400–2500 nm), light‐saturated photosynthesis (Asat), leaf dark respiration (Rdark), leaf mass area (LMA), and % leaf water across five campaigns in a six‐month period during which there were two causes of tree mortality: a major natural drought and a co‐incident tree stem girdling treatment. We find that prior to mortality, there were significant (p < 0.05) leaf spectral changes in the red (650–700 nm), the NIR (1,000–1,400 nm), and SWIR bands (2,000–2,400 nm) and significant reductions in the potential carbon balance of the leaves (increased Rdark and reduced Asat). We show that the partial least squares regression technique can predict mortality in tropical trees across different species and functional groups with medium precision but low accuracy (r2 of .65 and RMSE/mean of 0.58). However, most tree death in our study was due to girdling, which is not a natural form of death. More research is needed to determine if this spectroscopy technique can be applied to tropical forests in general.

Forest Structural Complexity and Biomass Predict First-Year Carbon Cycling Responses to Disturbance

The pre-disturbance vegetation characteristics that predict carbon (C) cycling responses to disturbance are not well known. To address this gap, we initiated the Forest Resilience Threshold Experiment, a manipulative study in which more than 3600 trees were stem girdled to achieve replicated factorial combinations of four levels (control, 45, 65, and 85% gross defoliation) of disturbance severity and two disturbance types (targeting upper or lower canopy strata). Applying a standardized stability framework in which initial C cycling resistance to disturbance was calculated as the first-year natural log response ratio of disturbance and control treatments, we investigated to what extent pre-disturbance levels of species diversity, aboveground woody biomass, leaf area index, and canopy rugosity—a measure of structural complexity—predict the initial responses of subcanopy light-saturated leaf CO2 assimilation (Asat), aboveground wood NPP (ANPPw), and soil respiration (Rs) to phloem-disrupting disturbance. In the year following stem girdling, we found that above-ground C cycling processes, Asat and ANPPw, were highly resistant to increases in disturbance severity, while Rs resistance declined as severity increased. Disturbance type had no effect on first-year resistance. Pre-disturbance aboveground woody biomass, and canopy rugosity were positive predictors of ANPPw resistance and, conversely, negatively related to Rs resistance. Subcanopy Asat resistance was not related to pre-disturbance vegetation characteristics. Stability of C uptake processes along with Rs declines suggest the net C sink was sustained in the initial months following disturbance. We conclude that biomass and complexity are significant, but not universal, predictors of initial C cycling resistance to disturbance. Moreover, our findings highlight the utility of standardized stability measures when comparing functional responses to disturbance.

Phenology and Dispersal of the Wheat Stem Sawfly (Hymenoptera: Cephidae) Into Winter Wheat Fields in Nebraska.

Historically, the wheat stem sawfly, Cephus cinctus Norton was a pest in spring wheat-growing regions of the northern Great Plains. However, in the 1980s, it was found infesting winter wheat fields in Montana. Infestations were first detected in western Nebraska in the 1990s, and have since spread throughout the Nebraska Panhandle. Larval damage occurs from stem-mining, but stem girdling that results in lodged stems that are not harvested results in the greatest yield losses. The biology and phenology of the wheat stem sawfly are well described in the northern portion of its range, but they are lacking in Colorado, southeast Wyoming, and Nebraska. In this study, the phenology and dispersal of the wheat stem sawfly in Nebraska winter wheat fields is described using sweep net and larval sampling. During this 2-yr study, adult activity began on May 23 and ended on June 21. Adult sex ratios were 2.32 males per female in 2014 and 0.46 males per female in 2015. Both sexes demonstrated an edge effect within the wheat fields, with greater densities near the field edge. The edge effect was stronger for male wheat stem sawfly than females. Wheat stem sawfly larval density also had an edge effect, regardless of the density of female wheat stem sawfly present. This information will be useful for developing management plans for the wheat stem sawfly in Nebraska and neighboring regions.

Study on the effects of polymer modifiers and phloem girdling on cotton in cadmium-contaminated soil in Xinjiang Province, China

The effects of two liquid modifiers (polyacrylate compound modifier and organic polymer compound modifier) and phloem girdling (stem girdling and branch girdling) on cadmium (Cd) content, Cd transport, and photosynthetic parameters of cotton (Xinluzao 60) in Cd-contaminated soil (40 mg kg −1) were studied through barrel experiment. The results showed that the distribution ratios of Cd in stem, leaves, and bolls, leaf net photosynthetic rate (Pn), leaf stomatal conductance (Gs), leaf transpiration rate (Tr), and chlorophyll content were decreased after girdling; and the application of modifiers reduced the Cd content and the Cd transported to the shoot, while alleviating photosynthetic damage caused by girdling. In general, our results indicated that the inhibition of carbohydrate supply caused by girdling reduced the photosynthetic capacity of cotton, while the applications of the two liquid modifiers decrease the influence to cotton photosynthesis. Moreover, Cd and modifiers may be transported to the shoot through both phloem and xylem.

Spatial and temporal dynamics of the fungal community of chestnut blight cankers on American chestnut (Castanea dentata) in Michigan and Wisconsin

Abstract Chestnut blight cankers caused by the fungal pathogen Cryphonectria parasitica on infected American chestnut trees can be invaded by hypoviruses that infect C. parasitica mycelia. Hypoviruses lower pathogen virulence (i.e. hypovirulence) and decrease the probability of stem girdling. Cankers also are susceptible to invasion by non-C. parasitica fungi; however, the influence of invading fungi on disease severity is unknown. Fungi may antagonize C. parasitica growth and combine with hypovirulence to further reduce the probability of stem girdling. We conducted a survey to investigate how spatial and temporal dynamics of the fungal community within cankers correlate with canker severity and the likelihood of girdling. Cankers from six American chestnut populations were monitored from 2012 to 2016. We found that the spatial distribution of the fungal community within a canker resembled a mosaic that increased in fungal diversity over time. Cankers with increased fungal diversity were not associated with higher stem survivorship over time, compared to cankers with abundant hypovirulence in recovering chestnut populations. Fungal communities also were unstable, and cankers on surviving trees containing hypovirulence consistently were invaded by non-C. parasitica fungi. Our results suggest that canker communities are not stable, and volatility of the fungal community indicates that cankers can change quickly from less severe to more severe cankers through loss of hypovirulence. Non-C. parasitica fungi may facilitate canker expansion through greater inhibition of hypovirulent C. parasitica relative to virulent C. parasitica, which may permit virulent C. parasitica to escape hypovirus infection and resume rapid canker expansion.

Hendrik Poorter.

Hendrik Poorter.

Effect of organic matter manipulation on the seasonal variations in microbial composition and enzyme activities in a subtropical forest of China

Purpose The aim of this study was to determine the impacts of reduced aboveground and belowground C inputs on the community composition of soil microbes and enzyme activities in a seasonal context. Materials and methods Litterfall removal, root exclusion, and stem girdling treatments under a subtropical conifer plantation growing on a coarse texture of sandy soil in southeast China were employed. One year after the initiation of the treatments, we measured the soil microbial biomass, community composition, and enzyme activities, including hydrolytic and oxidative extracellular enzymes on a seasonal basis. Soil inorganic N, dissolved organic C and N, and available P were also determined. Results and discussion Seasonal variations of soil microbial composition and enzyme activities were attributed to soil temperature and moisture and soil nutrient availability. Girdling treatments significantly increased the abundances of gram-negative bacteria and actinomycetes in winter when soil temperature, moisture, and available nutrients were at the lowest level among the four seasons. Girdling alone and girdling combined with litter removal and root trenching significantly decreased the cellobiohydrolase, β-glucosidase, β-1,4-N-acetylglucosaminidase, and acid phosphatase activities in autumn. These hydrolytic enzyme activities were significantly correlated with soil moisture, NH4+, DOC, and available P. We also found a significant relationship between hydrolytic enzyme activities and the ratio of gram-positive to gram-negative bacteria. Conclusions Plant belowground C allocation, soil temperature, and moisture drove the seasonal patterns of soil microbial composition and enzyme activities. Labile C input by root exudates is a key determinant of ecosystem functions mediated by soil microbes such as microbial decomposition processes.

Influence of girdling on flower sex ratio, biochemical constituents, and fruit set intensity in mango (Mangifera indica L.)

The study was conducted in mango to understand the relationship of various metabolites with flower sex ratio (hermaphrodite/male) and fruit set using stem girdling technique. Girdling one year old branches was carried out in two cultivars, Mangifera indica 'Alphonso' and 'Totapuri' to retain 50, 100 and 150 leaves to vary the content of hormones, sugars, total carbohydrates, and amino acids. The increased leaf number was found to increase the total carbohydrates, glucose, fructose, and other sugars. Girdling also significantly increased the concentration of abscisic acid (ABA), salicylic acid (SA), jasmonic acid (JA), indole acetic acid, indole butyric acid but decreased gibberellins and cytokinins. Amino acids also increased in the girdled branches. Girdling improved the flower sex ratio as well as fruit set more efficiently in 'Alphonso' than in 'Totapuri'. Results indicate that the increase in production of hermaphrodite flowers may be more related to the higher production of growth inhibitors, like ABA, SA, and JA along with increased concentration of sugars. Increased auxin concentration also might play an important role in increasing the fruit set.

Effects of girdling on stem CO2 efflux and its temperature sensitivity in Chinese fir and sweetgum trees

The effect of carbohydrate supply on stem CO2 efflux and its temperature sensitivity is crucial in predicting forest carbon cycling under climate change and forest management but has been rarely reported. In this study, we altered carbohydrate supply via stem girdling and investigated the effect of nonstructural carbohydrate concentration on stem CO2 efflux and its temperature sensitivity in Chinese fir and sweetgum trees in spring and winter. The stem CO2 efflux of Chinese fir increased significantly above the girdle in spring and winter, but decreased below the girdle in winter. The stem CO2 efflux of sweetgum increased above the girdle and decreased below the girdle in spring but not in winter. Seasonal variations in the response of stem CO2 efflux to girdling were associated with the changes in stem soluble sugar and starch concentration, especially in sweetgum tree. Girdling altered Q10 of the stem CO2 efflux, and this effect was dependent on location, season and species. Compared with the Q10 of the stem CO2 efflux in non-girdled trees, that of CO2 efflux in Chinese fir increased by 45% above the girdle but decreased by 16% below the girdle in winter. These values also exhibited 6% and 11% responses in spring. By contrast, the Q10 of the stem CO2 efflux in sweetgum increased by 95% above the girdle and decreased by 45% below the girdle in spring. However, these values were nearly constant in winter. We concluded that the effect of carbohydrate supply on stem CO2 efflux and its temperature sensitivity depends on season and species. Therefore, the vital role of carbohydrate supply that regulates stem CO2 efflux and its temperature sensitivity should be considered for the understanding and modeling of forest carbon cycling response to forest management, especially under global climate changes.

Stem girdling uncouples soybean stomatal conductance from leaf water potential by enhancing leaf xylem ABA concentration

To understand the impact of shoot-to-root ABA transport on water potential of, and xylem ABA concentration in, different plant tissues during soil drying, soybean (Glycine max cv. Siverka) plants were subjected to drought and girdling in a factorial experiment. Girdling was achieved by surgically excising the phloem tissue from just above the cotyledonary node. After girdling and withholding water, ABA concentrations were determined in xylem saps extracted from individual leaves, detached shoots and de-topped roots, after measuring stomatal conductance (gs), tissue water potentials, and root ABA concentrations. Soil drying decreased water potential throughout the plant and approximately doubled xylem ABA concentrations, coinciding with stomatal closure. Girdling slightly enhanced water potential, especially in droughted plants. Girdling diminished the soil-drying induced increase in xylem sap ABA concentration, and completely prevented root tissue ABA accumulation. Furthermore, girdling decreased root ABA concentration and increased leaf xylem ABA concentration of well-watered (WW) plants. Stomatal conductance declined linearly with leaf water potential only in intact plants, while gs declined as leaf xylem ABA concentration increased, independently of girdling. Thus shoot to root ABA transport not only determines (soil-drying induced) root ABA accumulation, but also limits ABA accumulation in the shoot to maintain stomatal opening of WW plants.

Long-term effects of stem girdling on needle structure in Scots pine

Abstract: Stem girdling is the process of completely removing a strip of cork and phloem tissue. Phloem is the living tissue which serves as the main long-distance pathway for transporting carbohydrates produced during photosynthesis to all parts of the plant where needed, from source leaves to sinks. Stem girdling has been used to study several functional aspects of phloem and their direct impacts on tree growth. Although both photosynthesis and transpiration processes take place in needles, no studies exist which investigate the effect of source-sink disturbance on needle structure. In this study, we evaluated changes in needle morphology and anatomy in current-year Scots pine needles 227 and 411 days after girdling (DAG). Although the studied needle parameters recorded 227 DAG were from 2 to 20% higher than the same parameters in control needles, the differences were not significant. On the other hand, needles 411 DAG were thinner, with decreased cross-sectional areas, phloem areas, vascular cylinder areas, needle dry mass, needle density, and needle flatness when compared to control needles. Marked variations in needle growth were observed 411 DAG, with a smaller number of correlations among almost all studied needle parameters in needles 411 DAG when compared to control needles or needles 227 DAG. Structural development determining needle flatness, needle density, and leaf mass per area (LMA) appeared to have driving factors that were independent of the other studied needle parameters, as correlations with other parameters were not significant in any treatment. The changes in overall needle structure observed after long-term stem girdling provide new insights into the processes that occur as a result of source-sink disturbances. This type of data could be helpful, for example, in studies specifically focused on phloem transport, tree carbon relationships, or investigations modeling gas exchange. Our study might also support gene expression studies, which could provide further knowledge about the regulatory mechanisms that determine needle size and structural form.

Living on borrowed time - Amazonian trees use decade-old storage carbon to survive for months after complete stem girdling.

Summary Nonstructural carbon (NSC) reserves act as buffers to sustain tree activity during periods when carbon (C) assimilation does not meet C demand, but little is known about their age and accessibility; we designed a controlled girdling experiment in the Amazon to study tree survival on NSC reserves.We used bomb‐radiocarbon (14C) to monitor the time elapsed between C fixation and release (‘age’ of substrates). We simultaneously monitored how the mobilization of reserve C affected δ13 CO 2.Six ungirdled control trees relied almost exclusively on recent assimilates throughout the 17 months of measurement. The Δ14C of CO 2 emitted from the six girdled stems increased significantly over time after girdling, indicating substantial remobilization of storage NSC fixed up to 13–14 yr previously. This remobilization was not accompanied by a consistent change in observed δ13 CO 2.These trees have access to storage pools integrating C accumulated over more than a decade. Remobilization follows a very clear reverse chronological mobilization with younger reserve pools being mobilized first. The lack of a shift in the δ13 CO 2 might indicate a constant contribution of starch hydrolysis to the soluble sugar pool even outside pronounced stress periods (regular mixing).

Reducing damping-off problems in eggplant (Solanum melongena L.): A participatory testing of nursery management in Bangladesh

Eggplant seedling production in homestead nurseries of farmers in Jamalpur (Bangladesh) is greatly compromised by damping-off. Therefore, farmers often do not have enough seedlings to transplant. Effective treatments of soil and seed to reduce disease pressure in the nurseries are available but little is known on the relative contributions of soil-borne and seed-borne pathogens to damping-off and on how nursery management can integrate management options under farmers' conditions. A 2-year nursery study was conducted in consecutive seasons, jointly with farmers and using farmers' preserved seed and farmers' nurseries subject to damping-off problems. Year 1 involved a single nursery, Year 2 nine nurseries. The following treatments were tested: Trichoderma harzianum as soil amendment combined with seed treatment using either hot water or Carbendazim and farmers' conventional practice: curative spraying after appearance of damping-off. In Year 1, a control (no intervention) treatment was also included. Emergence of seedlings, incidence of damping-off, seedling performance variables and farmers' appreciation of seedling vigour were recorded. Soil treatment with T. harzianum combined with seed treatment with hot water increased seedling emergence and produced 25–64 percentage points more healthy seedlings than farmers' conventional practice. This combined treatment also improved seedling performance (height, root length, lateral root development), and reduced stem girdling, a symptom associated with disease infection of transplants. To determine the contributions of nursery soil and farmers’ seed to damping-off, blotter and in vitro studies in the laboratory and tray studies in a screen house were performed. Soil proved to be the major pathogen source and treating nursery soil with T. harzianum had the largest positive effect on seedling performance, both reducing damping-off and enhancing seedling growth.

Lentil anthracnose: epidemiology, fungicide decision support system, resistance and pathogen races

Colletotrichum lentis causes anthracnose of lentil (Lens culinaris) in Canada that results in defoliation, stem girdling and severe yield losses, but the disease is rarely reported elsewhere. The pathogen survives as microsclerotia on lentil debris for up to 3 years when buried in the soil, but loses viability on the soil surface. Windborne debris spreads the pathogen to neighbouring fields, while seedborne infection is less important. Foliar fungicides were registered, and a fungicide decision support system was developed which assessed disease risk with 85% accuracy. Around 2300 L. culinaris accessions from 50 countries were screened for resistance. Congruently, two races – Ct1 and Ct0 – were identified on differential lentil lines. Resistant lines were generated by cycles of inoculation and selfing of single resistant plants which resulted in the following three accessions resistant to both races: VIR2633 (Georgia), VIR2058 and VIR2076 (Czech Republic), while six and 49 lines had resistance to Ct0 and Ct1, respectively. Ct1 resistance is controlled by recessive and dominant genes crt1 and CtR3 in variety ‘Indianhead’, ctr2 and CtR5 in accession PI345629 and CtR4 in PI320937. Molecular markers linked to Ct1 resistance were identified on linkage group six, close to Ascochyta lentis resistance, and were used to combine resistance to both pathogens in breeding lines. Two repeat rich regions in the intergenic spacer (IGS) of ribosomal DNA can be used to differentiate the two C. lentis races. Utilizing length polymorphisms in a 39 nucleotide repeat region showed the races were equally frequent among isolates collected between 1991 and 1999, while 95% belonged to race Ct0 in 2010, likely because lentil varieties are susceptible to race Ct0, but around one-third of the varieties had Ct1 resistance.

Phloem Girdling of Norway Spruce Alters Quantity and Quality of Wood Formation in Roots Particularly Under Drought.

Carbon (C) availability plays an essential role in tree growth and wood formation. We evaluated the hypothesis that a decrease in C availability (i) triggers mobilization of C reserves in the coarse roots of Picea abies to maintain growth and (ii) causes modification of wood structure notably under drought. The 6-year-old saplings were subjected to two levels of soil moisture (watered versus drought conditions) and root C status was manipulated by physically blocking phloem transport in the stem at three girdling dates (GDs). Stem girdling was done before the onset of bud break [day of the year (doy) 77], during vigorous aboveground shoot and radial stem growth (GD doy 138), and after cessation of shoot growth (GD doy 190). The effect of blockage of C transport on root growth, root phenology, and wood anatomical traits [cell lumen diameter (CLD) and cell wall thickness (CWT)] in earlywood (EW) and latewood (LW) was determined. To evaluate changes in belowground C status caused by girdling, non-structural carbohydrates (soluble sugars and starch) in coarse roots were determined at the time of girdling and after the growing season. Although fine root mass significantly decreased in response to blockage of phloem C transport, the phenology of root elongation growth was not affected. Surprisingly, radial root growth and CLD of EW tracheids in coarse roots were strikingly increased in drought-stressed trees, when girdling occurred before bud break or during aboveground stem growth. In watered trees, the growth response to girdling was less distinct, but the CWT of EW significantly increased. Starch reserves in the roots of girdled trees significantly decreased in both soil moisture treatments and at all GDs. We conclude that (i) radial growth and wood development in coarse roots of P. abies saplings are not only dependent on current photosynthates, and (ii) blockage of phloem transport induces physiological changes that outweigh drought effects imposed on root cambial activity and cell differentiation.

Moderate Disturbance Has Similar Effects on Production Regardless of Site Quality and Composition

Moderate severity disturbances, which only kill a subset of canopy trees (e.g., via insects, pathogens, and windthrow), are increasingly widespread in North America, and can alter forest structure and production. Whether the net primary production (NPP) of forest stands differing in pre-disturbance site quality and composition respond similarly to moderate severity disturbance, however, is unknown, but critical to understanding the disturbance response dynamics of patchy landscapes. We experimentally disturbed three, 2-ha stands varying in pre-disturbance primary production and community composition, temporarily reducing live stand basal area by 38% to 66% through the stem girdling of all mature early successional aspen (Populus tremuloides Michx. and Populus grandidentata Michx.) and birch (Betula papyrifera Marshall). Disturbance significantly altered stand-scale physical and biological structure and prompted a similar decade-long pattern of wood NPP decline and recovery. All stands exhibited an initial reduction in wood NPP, followed by a recovery period and eventual return to pre-disturbance levels within eight years, with the most productive stand exhibiting an increase in primary production following recovery. Following wood NPP recovery, more biologically diverse forest canopies with higher leaf area indexes intercepted more light, and, consequently, had higher rates of wood NPP. We conclude that, despite substantial pre-disturbance differences in productivity and community composition, relative wood NPP recovery patterns can be similar, though long-term post-recovery primary production may trend higher in more productive and compositionally diverse stands. We suggest that improved mechanistic understanding of different forest ecosystems’ responses to disturbances remains critical to informing management decisions across diverse landscape mosaics.

Stored root carbohydrates can maintain root respiration for extended periods.

Tight coupling between below-ground autotrophic respiration and the availability of recently assimilated carbon (C) has become a paradigm in the ecophysiological literature. Here, we show that stored carbohydrates can decouple respiration from assimilation for prolonged periods by mobilizing reserves from transport roots to absorptive roots. We permanently disrupted the below-ground transfer of recently assimilated C using stem girdling and root trenching and measured soil CO2 efflux for over 1 yr in longleaf pine (Pinus palustris), a species that has large reserves of stored carbohydrates in roots. Soil CO2 efflux was not influenced by girdling or trenching through the 14-month observation period. Stored carbohydrate concentrations in absorptive roots were not affected by the disrupted supply of current photosynthate for over 1 yr; however, carbohydrate concentrations in transport roots decreased. Our results indicate that root respiration can be decoupled from recent canopy assimilation and that stored carbohydrates can be mobilized from transport roots to absorptive roots to maintain respiration for over 1 yr. This refines the current paradigm that canopy assimilation and below-ground respiration are tightly coupled and provides evidence of the mechanism and dynamics responsible for decoupling the above- and below-ground processes.

How timing of stem girdling affects needle xylem structure in Scots pine

While needles represent a proportionally large fraction of whole-plant hydraulic resistance, no studies to date have investigated how source–sink disturbances affect needle xylem structure. In this study, we evaluated structural changes in xylem in current-year needles of Scots pine 227 and 411 days after stem girdling (hereafter referred to as DAG). Maximum and minimum tracheid lumen diameters and therefore also the size of tracheid lumen areas increased in needles 227 DAG compared to control needles. In contrast, tracheid dimensions were similar in needles 411 DAG as in the control needles, but smaller xylem area and lower number of tracheids resulted in the lower theoretical needle hydraulic conductivity of those needles. Several needle xylem parameters were intercorrelated in both control and girdled trees. These observed changes provide a new understanding of the processes that occur following a source–sink disturbance. Considering anatomical parameters such as the number of tracheids, tracheid dimension, or needle xylem area, which are rarely described in physiological studies, could be helpful, for example, in understanding to tree hydraulic systems or for modeling gas exchange. Finally, empirical equations were developed to calculate needle theoretical hydraulic conductivity and the number of tracheids in needles using an easily measurable parameter of needle xylem area.

The effects of burning season and severity on the mortality over time of Pinus nigra spp. salzmannii (Dunal) Franco and P. sylvestris L.

Abstract Understanding tree mortality after fires can be used to improve planning of prescribed burning and post-burning salvage logging. This study aims to assess the effects of burning season, fire severity and tree size on the evolution of post-burning mortality of pine species with different fire resistance traits. In 2013, two paired plots (1 ha each) were burned, one in spring and the other in fall, in two mixed-stands of P. sylvestris and P. nigra, and the fire severity and tree size variables were measured. Mortality was assessed individually in a total of 933 pines during the following 3 years post-burning. We modeled the initial (1 year post-burning) and delayed (3 years post-burning) tree mortality using Firth logistic regression and Conditional Inference trees. The probability of dying after burning was lower in P. nigra than in P. sylvestris confirming that P. nigra has higher fire-resistance compared to the moderate fire-resistance of P. sylvestris . Initial mortality was slightly higher for pines burned in spring than for those burned in fall; however, after 3 years the proportion of pines that had died after the fall burns was higher than that after the spring burns. Nevertheless, for the two periods modeled, the likelihood of a pine dying was significantly higher in spring than in fall given a certain level of crown scorch. This suggests that pines with damaged crowns were more likely to die when burning occurred during periods of active growth. However, the higher fire severities at the base of the trees that occurred during the fall burns compared to the spring burns are probably the cause of the higher rates of delayed tree mortality. Higher likelihood of mortality in trees burned in fall, with higher percentages of white ashes in the stem base suggest basal stem girdling as a cause of delayed pine mortality. We can conclude that crown damage is more detrimental when burning occurs during the growth season, while longer combustion times during fall burns may have caused the higher delayed pine mortality in comparison with spring burns.