Effects of xylem cavitation and freezing injury on dieback of yellow birch (Betula alleghaniensis) in relation to a simulated winter thaw

Shoot dieback, shoot growth, stem xylem cavitation, stem and root freezing injury, and root pressure were measured in 2-year-old, cold-hardened, potted yellow birch (Betula alleghaniensis Britt.) seedlings that had been subjected to a simulated winter thaw for 0, 5, 10, 19 or 27 days followed by 10 weeks at -10 degrees C. Stem xylem cavitation was determined as percent loss of hydraulic conductivity. Stem freezing injury was measured as electrolyte leakage (EL). Root freezing injury was determined by EL and by triphenyl tetrazolium chloride (TTC) reduction. Thaw duration was significantly correlated with dieback, new shoot growth, stem xylem cavitation, stem and root freezing damage, and root pressure (P < 0.05). In particular, shoot dieback was positively correlated with stem xylem cavitation (P < 0.001), residual stem xylem cavitation (P < 0.01) and root freezing injury (P < 0.010), but only weakly correlated with stem freezing damage (P < 0.05). In roots, freezing damage was negatively correlated with root pressure (P < 0.05), which, in turn, was negatively correlated with residual stem xylem cavitation after root pressure development. In stems, there was no correlation between freezing damage and xylem cavitation. We conclude that long periods of winter thaw followed by freezing resulted in freezing injury to roots concomitant with a reduction in root pressures, leading to poor recovery from freezing-induced xylem embolism.     

Effects of simulated thaw on xylem cavitation,residual embolism,spring dieback and shoot growth in yellow birch

Yellow birch seedlings (Betula alleghaniensis Britt.) that had lost more than 90% of their stem hydraulic conductivity during ambient winter temperatures were exposed to 0 and 20 days of a simulated winter thaw followed by a 48-h freezing treatment at 0, -5, -10, -20 and -30 degrees C. After measuring freezing injury to shoots and roots, the seedlings were placed in a greenhouse where recovery of xylem conductivity and new growth were measured. Shoot xylem cavitation was measured as percent loss of hydraulic conductivity. Shoot freezing injury was assessed by electrolyte leakage (EL) and root freezing injury was assessed by EL and triphenyl tetrazolium chloride reduction. Seedlings pretreated with thaw had higher stem water contents and suffered more freezing damage to roots and shoots (at -20 and -30 degrees C, respectively) than unthawed seedlings. After 3 weeks in a greenhouse, seedlings from the 0, -5 and -10 degrees C freezing treatments showed complete recovery of xylem conductivity, with substantially increased stem water contents. Poor recovery of hydraulic conductivity was observed only in seedlings that were subjected to freezing treatments at -20 and -30 degrees C, regardless of thaw treatment. Of these embolized seedlings, however, only those not previously thawed showed recovery of hydraulic conductivity or regained stem water content after 9 weeks in the greenhouse. Shoot dieback, bud burst and length of new shoots were significantly related to the extent of stem xylem cavitation and freezing injury. We conclude that (1) the simulated winter thaw predisposed yellow birch seedlings to freezing damage in shoots and roots by dehardening tissues and increasing their water content; (2) root freezing damage in turn affected the seedlings' ability to refill embolized stem xylem, resulting in considerable residual xylem embolism after spring refilling; (3) further recovery of stem xylem conductivity was attributable to growth of new vessels; (4) and the permanent residual embolism, together with root and shoot freezing injury, caused increased dieback, bud mortality and reduced growth of new shoots.     

Responses of xylem cavitation,freezing injury and shoot dieback to a simulated winter thaw in yellow birch seedlings growing in different nursery culture regimes

Seedlings of yellow birch originating from the same seed source were treated with two levels of fertilizers during two growing seasons. The lower level of fertilizers, such as 50 (11:41:08 N:P:K ratio), 100 (20:08:20) and 35 ppm of nitrogen (08:20:30) were applied as the starter, grower and finisher, respectively. The higher level ones consisted of 75, 150, and 100 ppm of nitrogen in the same fertilizers. After 2 years growth, seedlings treated with the higher level of fertilizers, had fewer lateral branches, greater height and larger stem diameter. After natural hardening from November to February, seedlings were subjected to 0, 5, 10, 19 and 27 days of a simulated winter thaw followed by 10 weeks at −10°C. After the thaw-refreeze treatments, series of measurements were carried out. Stem xylem cavitation and root freezing injury significantly increased with thaw duration regardless of levels of fertilizer treatments. Stem freezing injury also significantly increased with thaw duration in all stem segments of seedlings treated with the higher level of fertilizers, but only in the top segments for the seedlings provided with the lower level of fertilizers. However, seedlings treated with the higher level of fertilizers developed cavitation more quickly. After 1 month of growth in the greenhouse following the thaw and freeze treatment, both types of seedlings showed significant decrease in the length of new shoot growth and increase in percent length of shoot dieback with thaw duration. The length of new shoots, however, was always greater for the plants treated with the higher level of fertilizers. No difference of shoot dieback between the seedlings of the two different nursery treatments were observed. Correlation analyses showed that the length of new shoots was highly related to root and stem freezing injury, while dieback was best correlated with root freezing injury and stem xylem cavitation regardless of the levels of fertilizer treatments. It was concluded that (1) the higher level of fertilizer applied during the culture of yellow birch seedlings can accelerate xylem cavitation and dehardening in the stem following freeze–thaw events; (2) stem xylem cavitation was unlikely the cause of stem freezing injury; and (3) root freezing injury and stem xylem cavitation are the most reliable measurements for predicting dieback of potential planting stock, but both root and shoot freezing injury are relate well to regrowth of new shoots in stock exposed to prolonged thaw.     

Seasonal variation in xylem pressure of walnut trees: root and stem pressures

Measurements of air and soil temperatures and xylem pressure were made on 17-year-old orchard trees and on 5-year-old potted trees of walnut (Juglans regia L.). Cooling chambers were used to determine the relationships between temperature and sugar concentration ([glucose] + [fructose] + [sucrose], GFS) and seasonal changes in xylem pressure development. Pressure transducers were attached to twigs of intact plants, root stumps and excised shoots while the potted trees were subjected to various temperature regimes in autumn, winter and spring. Osmolarity and GFS of the xylem sap (apoplast) were measured before and after cooling or warming treatments. In autumn and spring, xylem pressures of up to 160 kPa were closely correlated with soil temperature but were not correlated with GFS in xylem sap. High root pressures were associated with uptake of mineral nutrients from soil, especially nitrate. In autumn and spring, xylem pressures were detected in root stumps as well as in intact plants, but not in excised stems. In contrast, in winter, 83% of the xylem sap osmolarity in both excised stems and intact plants could be accounted for by GFS, and both GFS and osmolarity were inversely proportional to temperature. Plants kept at 1.5 degrees C developed positive xylem pressures up to 35 kPa, xylem sap osmolarities up to 260 mosmol l(-1) and GFS concentrations up to 70 g l(-1). Autumn and spring xylem pressures, which appeared to be of root origin, were about 55% of the theoretical pressures predicted by osmolarity of the xylem sap. In contrast, winter pressures appeared to be of stem origin and were only 7% of the theoretical pressures, perhaps because of a lower stem water content during winter.     

Stem radial CO_2 conductance affects stem respiratory CO_2 fluxes in ash and birch trees

The CO_2 released from respiring cells in woody tissues of trees can contribute to one of three fluxes:efflux to the atmosphere(E_A),internal xylem sap transport flux(F_T),and storage flux(DS).Adding those fluxes together provides an estimate of actual stem respiration(R_S).We know that the relative proportion of CO_2 in those fluxes varies greatly among tree species,but we do not yet have a clear understanding of the causes for this variation.One possible explanation is that species differ in stem radial CO_2 conductance(g_c).A high g_c would favor the E_A pathway and a low g_cwould favor the F_Tpathway.However,g_chas only been measured once in situ and only in a single tree species.We measured g_cusing two methods in stems of Fraxinus mandshurica Rupr.(ash)and Betula platyphylla Suk.(birch)trees in situ,along with R_S,E_A,F_T and DS.Stem radial CO_2 conductance was substantially greater in ash trees than in birch trees.Corresponding to that finding,in ash trees over 24 h,E_Aconstituted the entire flux of respired CO_2 ,and F_Twas negative,indicating that additional CO_2 ,probably transported from the root system via the xylem,was also diffusing into the atmosphere.In ash trees,F_T was negative over the entire 24 h,and this study represents the first time that has been reported.The addition of xylem-transported CO_2 to E_A caused E_Ato be 9% higher than the actual R_Sover the diel measurement period.Birch trees,which had lower g_c,also had a more commonly seen pattern,with E_A accounting for about 80% of the CO_2 released from local cell respiration and F_T accounting for the remainder.The inorganic carbon concentration in xylem sap was also lower in ash trees than in birch trees:2.7 versus 5.3 mmol L^-1,respectively.Our results indicate that stem CO_2 conductance could be a very useful measurement to help explain differences among species in the proportion of respired CO_2 that remains in the xylem or diffuses into the atmosphere.     

Freezing cycles enhance winter injury in Picea rubens

We examined changes in chlorophyll absorbency in red spruce (Picea rubens Sarg.) foliage in response to simulated freezing cycles. Current-year branch tips were collected from 16 trees on January 8, January 20, February 8 and February 26, 1996. Tissue was subjected to freezing cycle treatments with a minimum of -35 degrees C and a maximum of 3 degrees C for a one-cycle treatment, and -9, -6, -3, 0 or 3 degrees C for four-cycle treatments. Samples were frozen at a rate of 5 degrees C h(-1), and warmed at 12 to 15 degrees C h(-1). Controls were held at -9 degrees C. Temperatures during the three-day periods preceding each sample date averaged -18, 4.7, -9.6 and 3.7 degrees C, respectively. On January 8, treated trees showed no significant (P > 0.1) increase in the breakdown of chlorophyll, as measured by the ratio of chlorophyll a absorbency (435 nm) to phaeophytin a absorbency (415 nm), compared with control branch tips. On later sampling dates, seven trees consistently exhibited needle reddening and nine exhibited few symptoms (< 10% of total needle surface reddened) after four-cycle treatments. On February 26, chlorophyll degradation in trees with needle reddening differed (P < 0.05) from the control by 26, 26, 16, 14 and 15% for the 3, 0, -3, -6 and -9 degrees C maxima, respectively. No detectable chlorophyll degradation occurred after a one-cycle treatment in any trees on any sampling date. Freezing cycles with sub-zero maxima and a -35 degrees C minimum enhanced winter injury in red spruce after a midwinter thaw had rendered the trees susceptible to freezing damage.     

Xyleborus glabratus attacks and systemic colonization by Raffaelea lauricola associated with dieback of Cinnamomum camphora in the southeastern United States

Laurel wilt, caused by Raffaelea lauricola, is responsible for extensive mortality of redbay and other American members of the Lauraceae in the southeastern United States. Raffaelea lauricola is a mycangial symbiont of the redbay ambrosia beetle (Xyleborus glabratus), and the beetle and fungus were accidentally introduced from Asia. Branch dieback of camphortree (Cinnamomum camphora), an Asian member of the Lauraceae, has been occasionally observed in areas where laurel wilt has decimated redbay populations, and R. lauricola was isolated from such camphortrees. However, the role of X. glabratus and R. lauricola in this branch dieback remains unclear. Examination of camphortrees on Jekyll Island, Georgia showed that healthy‐appearing trees and those with branch dieback had been attacked by X. glabratus, but the trees with branch dieback had four times as many beetle attacks. Raffaelea lauricola was routinely isolated from discoloured xylem near beetle tunnels in healthy trees and those with dieback. Single‐point inoculations with R. lauricola on stems of mature, healthy camphortree trees failed to induce wilt‐like symptoms or branch dieback, although areas of discoloration were scattered throughout the xylem, and R. lauricola was reisolated irregularly at various heights in some inoculated trees. In growth chamber experiments, single‐point inoculations with R. lauricola resulted in systemic colonization but no wilt symptoms or branch dieback in camphortree saplings. In contrast, inoculations at multiple points along the stem (simulating multiple attacks by the vector) caused branch dieback and wilt‐like symptoms, including a brownish, diffuse discoloration of the xylem. Camphortree appears to be more resistant than American species of Lauraceae to the vascular wilt caused by R. lauricola. The fungus does colonize camphortrees systemically, however, and can apparently cause branch dieback. This suggests that the fungus may provide brood material for X. glabratus in Asia as it does in the southeastern United States.     

Xylem cavitation in roots and stems of Douglas-fir and white fir

Roots of hardwoods have been shown to be more vulnerable to xylem cavitation than stems. This study examined whether this pattern is also observed in a conifer species. Vulnerability to cavitation was determined from the pressure required to inject air into the vascular system of hydrated roots and stems, and reduce hydraulic conductance of the xylem. According to the air-seeding hypothesis for the cavitation mechanism, these air pressures predict the negative xylem pressure causing cavitation in dehydrating stems. This was evaluated for stems of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and white fir (Abies concolor (Gord. & Glend.) Lindl.). The air-injection method was applied to roots and stems of different sizes and positions in Douglas-fir trees. Roots, especially smaller roots with a xylem diameter < 5 mm, were more vulnerable to cavitation than stems. Mean cavitation pressure for smaller roots was -2.09 +/- 0.42 versus -3.80 +/- 0.19 MPa for larger roots (> 8 mm diameter). Within the shoot system, smaller stems (< 5 mm diameter) were most vulnerable to cavitation, having a mean cavitation pressure of -4.23 +/- 0.565 versus -5.27 +/- 0.513 MPa for large stems (> 8 mm diameter). There was no correlation between tracheid diameter and mean cavitation pressure within root or stem systems, despite larger tracheid diameters in roots (23.3 +/- 3.9 micro m) than in stems (9.2 +/- 1.6 micro m). Smaller safety margins from cavitation in roots may be beneficial in limiting water use during mild drought, and in protecting the stem from low xylem pressures during extreme drought.     

Competition as a predisposing factor of crown dieback in a secondary forest of <Emphasis Type="Italic">Betula maximowicziana</Emphasis> in Hokkaido,northern Japan

We analyzed the probability that Betula maximowicziana Regel (monarch birch) would suffer crown dieback (crown-dieback probability) and the basal area growth rate (GB), which was found to be a predisposing stress factor making birch trees susceptible to crown dieback. First, we analyzed the relationship between the probability that birch trees would suffer from crown dieback in 1999 and GB from a period prior to the occurrence of crown dieback (1985–1987), using a data set of repeated measurements on 217 trees. Logistic regression analysis revealed that monarch birch had a larger crown-dieback probability when GB was low in the preceding period. Hence, there were predisposing stress factors that reduced GB and continued to affect trees for at least a decade. Next, we analyzed GB in the same period in relation to symmetrical and asymmetrical competition between trees and found that GB was reduced by symmetrical competition, suggesting that this was one of the predisposing factors for crown dieback. Based on these results, we used selected models for crown-dieback probability and GB to calculate crown-dieback probabilities for individuals with different initial basal areas and experiencing different intensities of symmetrical competition. The predicted crown-dieback probability decreased with decreasing symmetrical competition between trees. We discuss a possible process of crown dieback to death for monarch birch and the use of thinning as a method to reduce the risk of crown dieback.     

Integration of water transport pathways in a maple tree: responses of sap flow to branch severing

? It has been known for a long time that sectored and integrated patterns of vascular systems exist in different species and even within the same tree, depending on its age and history. However, very few publications consider the topology of the vascular pathways between roots and branches.

? Some results on this important aspect of the vascular system are presented in this paper. They have been obtained with adult maple trees by directly studying the water movement in the stem and root xylem with the heat field deformation (HFD) method for sap flow measurements.

? Multi-point HFD sensors were installed at different heights of a Norway maple tree (Acer platanoides L.) along its stem axis. Single-point HFD sensors were installed in three small lateral roots of another sample maple. Experimental treatments (branch severing) triggered changes in sap movement in the stem and root sapwood.

? The sample trees belong to the group with an integrated transport system (“integrated pipes”), sharing stem space on both sides of the tree to supply two large parts of the crown with water from each root sector. Nevertheless, conducting pathways had their autonomy for axial transport and the pipe model theory describes the vascular system of the studied trees well. Thus, the integration of axial transport in the stem xylem should presumably occur through the cross-grained network of axial vessels.

     

Winter variation in xylem sap pH of walnut trees: involvement of plasma membrane H+-ATPase of vessel-associated cells

We studied seasonal variation in xylem sap pH of Juglans regia L. Our main objectives were to (1) test the effect of temperature on seasonal changes in xylem sap pH and (2) study the involvement of plasma membrane H+-ATPase of vessel-associated cells in the control of sap pH. For this purpose, orchard-grown trees were compared with trees grown in a heated (> or = 15 degrees C) greenhouse. During autumn, sap pH was not directly influenced by temperature. A seasonal change in H+-ATPase activity resulting from seasonal variation in the amount of protein was measured in orchard-grown trees, whereas no significant seasonal changes were recorded in greenhouse-grown trees. Our data suggest that H+-ATPase does not regulate xylem sap pH directly by donating protons to the xylem, but by facilitating secondary active H+/sugar transport, among other mechanisms.     

Pre-commercial thinning,birch admixture and sprout management in planted Norway spruce stands in South Sweden

Early management of the regenerated seedlings shapes the future stand properties. To address these issues, pre-commercial thinning (PCT) and control treatments were applied to planted Norway spruce (Picea abies L. Karst) and naturally regenerated birch (Betula pendula Roth., Betula pubescens Ehrh.) stands in forest experiments in southern Sweden (lat. 56–57?N) containing 1.1–5.5?m tall saplings. The treatments were retention of 1000 or 2000?stems?ha^?1 of Norway spruce, with no birch or birch at 1000?stems?ha^?1. Treatments were replicated with and without annual removal of birch sprouts from stumps. The periodic annual increment (PAI) over five years was calculated for total stand volume and individual trees. The mean PAI of dominant trees was significantly higher both following all PCT treatments than controls, and following low rather than high-density PCT. Birch retention did not affect growth of the dominant trees but PAI was lower in plots with uncontrolled sprouting. The PAI of birch was significantly higher in low-density Norway spruce plots than in control plots and the high-density plots. The treatment response was significant even in stands with initial heights of only 1–2?m.     

Foliar Responses of Understorey Abies lasiocarpa to Different Degrees of Release Cutting in a Betula papyrifera and Conifer Mixed Species Stand

Foliar responses of subalpine fir [Abies lasiocarpa (Hook.) Nutt.] to thinning were studied in a 35-yr-old mixed stand of paper birch (Betula papyrifera Marsh.) and conifers. The stand regenerated naturally after a wildfire with a canopy dominated by paper birch (average height 9.8 m) and an understorey dominated by subalpine fir (average height 1.6 m). The stand was thinned to four densities of birch: 0, 600 and 1200 stems ha-¹ and control (unthinned at 2300-6400 stems ha-¹) in the autumn of 1995. The understorey conifers, mainly subalpine fir, were thinned to 1200 stems ha-¹. The study used a completely randomized split-plot design. Three sample trees were systematically selected from each treatment replicate and each tree stratum (upper, intermediate and lower understorey). One-year-old and older age class needles were collected from one south-facing branch within the fifth whorl from the tree top. Thinning of paper birch significantly (p <0.001) increased leaf area and dry weight per 100 needles for intermediate and short trees except in the 0 birch treatment. Understorey subalpine fir trees in 600 stems ha¹ birch (T3) had the largest leaf area and leaf dry weight per 100 1-yr-old needles. Specific leaf area (SLA) decreased from unthinned (T1) to 0 birch (T4). Lower understorey trees had the largest SLA. One-year-old needles had significantly higher N, P and K concentrations in all the thinning treatments. These responses are consistent with the shade tolerance of subalpine fir. The results suggest that when managing a paper birch-conifers mixed-wood forest it may be of benefit to understorey conifers to leave a birch canopy as a nursing crop.     

Photosynthetic capacity of red spruce during winter

We measured the photosynthetic capacity (P(max)) of plantation-grown red spruce (Picea rubens Sarg.) during two winter seasons (1993-94 and 1994-95) and monitored field photosynthesis of these trees during one winter (1993-94). We also measured P(max) for mature montane trees from January through May 1995. Changes in P(max) and field photosynthesis closely paralleled seasonal changes in outdoor air temperature. However, during thaw periods, field photosynthesis was closely correlated with multiple-day temperature regimes, whereas P(max) was closely correlated with single-day fluctuations in temperature. There was a strong association between short-term changes in ambient temperature and P(max) during the extended thaw of January 1995. Significant increases in P(max) occurred within two days of the start of this thaw. Repeated measurements of cut shoots kept indoors indicated that temperature-induced increases in P(max) can occur within 3 h. Although significant correlations between P(max) and stomatal conductance (g(s)) or intracellular CO(2) concentration (C(i)) raised the possibility that increases in P(max) resulted from increases in stomatal aperture, fluctuations in g(s) or C(i) explained little of the overall variation in P(max). Following both natural and simulated thaws, P(max) increased considerably but plateaued at only 37% of the mean photosynthetic rate reported for red spruce during the growing season. Thus, even though shoots were provided with near-optimal environmental conditions, and despite thaw-induced changes in physiology, significant limitations to winter photosynthesis remained.     

Biomass production and allometric above- and below-ground relations for young birch stands planted at four spacings on abandoned farmland

The objective of the study was to quantify above- and below-stumpbiomass of silver (Betula pendula Roth) and downy (Betula pubescensEhrh.) birches planted at four spacing intervals and growingon two soil types on an area of farmland. The 12-year-old bircheshad been grown at four spacings (1.3, 1.5, 1.8 and 2.6 m) ontwo sites: one on medium clay soil and the other on fine sandsoil. The dry weight of the stem, branches, leaves, stumps androots was estimated by drying and weighing sub-samples. Theprojected leaf area (PLA) m^–2 of trees, leaf area indexof stands and basic density (kg m^–3)of stems were alsoestimated. A significant greater dry weight of stem, branches,stump and roots and species and spacing for pendula birch werefound. The root length of silver birch was significantly greaterthan that for downy birch and for both species the root lengthwas greatest at the widest spacing (2.6 m). There was also asignificant difference between leaf weights of birch of thesame species growing on the two soil types. Significant differenceswere also found between PLA and species, and for both species,between PLA spacing. Basic density of stems was significantlydifferent between soil types. Equations for estimating the above-groundbiomass and root biomass from diameter at breast height weredeveloped for birches growing on fine sand and on medium claysoils. The total biomass production per hectare on fine sandwas higher for silver birch (19.9–65.9 tonnes ha^–1),than for downy birch (13.0–48.3 tonnes ha^–1). Onmedium clay soil, total biomass production for silver and downybirches was 30.8–52.8 and 16.8–42.8 tonnes ha^–1,respectively.     

Effects of thinning paper birch on conifer productivity and understory plant diversity

Naturally regenerated paper birch (Betula papyrifera Marsh.) is commonly removed from juvenile interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) plantations in southern interior British Columbia, Canada, to increase conifer productivity and create a free-growing stand; however, this practice is expensive and contentious because of possible negative ecological impacts. One solution is to retain an optimal density of birch where growth gains of understory Douglas-fir are balanced against losses to Armillaria ostoyae (Romagn.) Herink and understory plant species diversity. We sought to find this optimal density by comparing four evenly applied birch density reduction treatments (0, 400, 1111, and 4444 retained birch stems ha^?1) and an unthinned control (>7300 retained birch stems ha^?1). The mortality rate of Douglas-fir due to Armillaria root disease increased non-significantly with thinning intensity. Mean diameter increment of surviving Douglas-fir improved the most where birch was completely removed, with little variation among intermediate thinning treatments. Height growth was unaffected by the thinning treatments. Diversity of cryptogams was significantly greater in the control than where all birch was removed. We suggest that the treatment with 4444 retained birch stems ha^?1 provides the best balance for improving Douglas-fir growth while minimizing risk of increased Armillaria root disease and reduced understory plant diversity in young mixed stands.     

Effect of variable retention cutting on the relationship between growth of coarse roots and stem of Picea mariana

Silviculture heading for structural heterogeneity creates many single trees standing at stand margins, inner edges or in remnant tree groups. As they played just a minor role in the age class forest, the growth behaviour of strongly released trees is rather unexplored. Here we show how retention cutting, presently spreading in the boreal of Québec province, affects stem and coarse root growth of remained single black spruce [Picea mariana (Mill.) Britton]. Increment cores from roots and stems of 125 trees show that retention cutting triggers coarse root growth of the remaining trees. Compared with reference trees retention trees accelerate root in relation to stem growth. Mean and variability of the root–stem allometry significantly rise after retention cutting. The found acceleration of root in relation to stem growth means mechanical stabilisation of the retention trees and corroborates the retention cutting method. Evaluation of silvicultural treatments can be incomplete and misleading as long as they are just based on aboveground reactions and neglect root growth.     

Trichoderma atroviride promotes growth and enhances systemic resistance to Diplodia pinea in radiata pine (Pinus radiata) seedlings

Root drench application of Trichoderma atroviride isolates R32, R33, R40 and R84 promoted the growth of potted radiata pine seedlings. After 6 weeks, seedlings treated with R33 and R84 had thicker stems and greater stem and root biomass (p < 0.05) than untreated controls. Treatment with R32 increased seedling root biomass whilst R40 increased stem diameter. None of the isolates affected seedling height. One isolate, R33, induced systemic resistance to stem inoculation with Diplodia pinea and reduced dieback incidence by 20% compared with untreated controls. To our knowledge, this is the first report of systemic induced resistance by Trichoderma in a pine species. Furthermore, seedlings that were treated with R33 (root drench) plus foliar application of methyl jasmonate (MeJA) expressed elevated peroxidase activity in their stems 2 weeks later, compared with seedlings treated only with MeJA. Because R33 itself did not affect peroxidase activity, this may be indicative of treatment synergy or defence potentiation by R33. Curiously, R33 + MeJA induced terpenoids but suppressed phenylalanine ammonia‐lyase activity suggesting possible trade‐offs between phenolic and terpenoid defence pathways in the treated seedlings.     

Experimental evidence supporting the concept of light-mediated modulation of stem hydraulic conductance

It is a well-described phenomenon that plant leaves respond to changes in light intensity and duration by adjusting leaf hydraulic efficiency, and there is current consensus that up- or down-regulation of water channels (aquaporins) in the plasma membrane of the bundle sheath and mesophyll cells play a central role in the underlying mechanisms. Recently, experimental evidence has been provided also for light-mediated changes of stem hydraulic conductance (K(stem)) in field-grown laurel plants. This effect was attributed to differences in potassium ion concentration of xylem sap as a function of light conditions. In the present article, we report evidence obtained in silver birch (Betula pendula Roth), supporting the concept of light-mediated modulation of K(stem). Both canopy position (long-term effect) and current photosynthetic photon flux density (PPFD; short-term effect) had a significant impact (P < 0.001) on K(stem) measured in shoots taken from the lower (shade shoots) and upper (sun shoots) third of the crowns of ～25-year-old trees growing in a natural forest stand. The shade shoots responded more sensitively to light manipulation: K(stem) increased by 51% in shade shoots and 26% in sun shoots when PPFD increased from 70 to 330 μmol m?2 s?1. In 4-year-old trees growing in a dense experimental plantation, K(stem), specific conductivity of branch-wood (k(bw)) and potassium ion concentration ([K(+)]) in xylem sap varied in accordance with canopy position (P < 0.001). Both K(stem) and k(bw) increased considerably with light availability, increasing within the tree crowns from bottom to top; there was a strong relationship between mean values of K(stem) and [K(+)] in hydraulically sampled branches.     

Pilot study on the effects of elevated air temperature and CO<Subscript>2</Subscript> on artificially defoliated silver birch saplings

The impacts of elevated temperature and CO₂ on young silver birch (Betula pendula Roth) saplings after 0, 25, 50 or 75% artificial defoliation were assessed by measuring plant height and dry mass of aboveground compartments and roots and various morphological and physiological variables. Defoliation either increased or decreased plant growth depending on the severity of damage and the climatic treatment. At 21 °C and 400 mg L^?1 CO₂, defoliated plants were not able to compensate for the lost foliage, but growth compensation and adaptation to the changed conditions were greater; growth of young defoliated silver birch saplings increased, which led to increased height and a tendency to enhance final aboveground and root biomass and leaf nitrogen and carbon content compared to the nondefoliated controls. Nevertheless, the short-term effect of the different climatic conditions did not result in a significant overgrowth of defoliated plants. A slight increase in temperature and CO₂ were the most acceptable conditions for defoliated plants; however, a 4 °C increase with correspondingly higher CO₂ was more stressful as shown by less growth in height and biomass allocation to leaves, stems and roots. The findings from the pilot experiment are more applicable to young birch trees, but stress on young trees may be reflected in future tree growth.