Leaf and root growth dynamics in Eucalyptus globulus seedlings grown in drying soil

Summary Seedlings of Eucalyptus globulus growing in soil columns were subjected to a 24 day soil drying treatment. Water and solute potentials of both young expanding and fully expanded leaves declined under reduced soil water availability, while slightly higher turgor was sustained by the fully expanded leaves. Although leaf area of unwatered seedlings was smaller, the corresponding leaf dry weight was quite similar to that of well-watered seedlings. Soon after rewatering, leaf area of plants experiencing water shortage was comparable to that of well-watered plants. It seems that a difference in wall properties between juvenile and mature leaves allows for an effective pattern of water use by eucalypt plants growing in drying soil. Some stomatal opening is sustained and therefore, presumably, some carbon may be fixed, keeping the carbon balance of the whole plant positive, and allowing a continuous cell division despite the limited water supply. The highest root density of both well-watered and unwatered plants was found in the upper soil layers. However, root growth of unwatered seedlings was gradually increased in the deeper soil layers, where thicker root apices and higher soil water depletion rates per unit root length were recorded. As a consequence, root absorbing surface area was as large in unwatered plants as in well-watered plants.     

Detection and stability of quantitative trait loci (QTL) in Eucalyptus globulus

Eucalyptus globulus Labill. ssp. globulus is an important tree species for the pulp and paper industry, and several breeding programmes throughout the world are striving to improve key traits such as growth and wood density. This study aimed to detect quantitative trait loci (QTL) for growth, wood density, relative bark thickness and early flowering in a single full-sib E. globulus family grown across seven sites. Growth was measured a number of times over a 6-year period, enabling temporal stability of growth QTL to be studied. Ten putative QTL (LOD > 2.0) were detected in the single family, which was of moderate size. Based on permutations of the trait data, six of these QTL were significant at the experimentwise significance level of 0.1 for at least one of the four models implemented in analysis to remove site effects. For wood density, two putative QTL explained 20% of the variance for the trait, indicating that a small number of QTL might explain a reasonable proportion of the trait variance. One of these QTL was found to be independent of QTL for growth whereas the second QTL co-segregated with a QTL for relative incremental growth. The marker nearest to this QTL was associated with fast growth but low wood density. A putative growth QTL at year 6 was found to be relatively stable across ages. In addition, it was found that residuals from models based on measurements from across all families across all sites in the trial detected QTL with greater experimentwise significance.     

Diagnosis of nitrogen deficiency and toxicity of Eucalyptus globulus seedlings by foliar analysis

The relationship between shoot growth and foliar nitrogen (N) in E. globulus seedlings was studied in the glasshouse to determine standard values for N deficiency and toxicity diagnosis. Seedlings were grown for 9 weeks in yellow sand, at 10 rates of N, applied as ammonium sulphate, calcium nitrate or ammonium nitrate. Shoot dry weight (DW) increased linearly with N rate for all forms of N in the deficiency range. Seedlings continued to respond to higher rates of ammonium and ammonium nitrate than to nitrate. Maximum shoot DW for nitrate fed plants and ammonium nitrate fed plants were 51% and 84% respectively of ammonium fed plants. Total N concentration in the youngest fully expanded leaf (YFEL) ranged from 1.0% to 3.3% in deficient and adequate plants. The critical N concentration for deficiency diagnosis (corresponding to 90% maximum yield) in the YFEL, determined from these growth response curves averaged over all N forms, was 2.6% N. For ammonium nitrate fed plants, total N concentration in the YFEL for the severely deficient, deficient, adequate, and toxic ranges were <1.4%, 1.4–2.5%, 2.6–3.5%, > 4.3%. High total N concentrations were associated with growth depression and toxicity symptoms, which differed with N form. For nitrate fed plants, a total N concentration above 3.3% in the YFEL was associated with severe growth depression, and leaf tip necrosis. The adequate concentration range for ammonium nitrate was similar to values found on a field trial with 7 month old E. globulus trees grown on an exforest site.     

Impact of a eucalyptus (Eucalyptus globulus Labill.) plantation on the nutrient content and dynamics of coarse particulate organic matter (CPOM) in a small stream

Litterfall inputs, benthic storage and the transport of coarse particulate organic matter (CPOM) were studied in two headwater streams, one flowing through a mixed deciduous forest and one through a plantation of Eucalyptus globulus. Vertical and lateral traps, transported CPOM and benthic CPOM were sampled monthly to biweekly and sorted into four categories: leaves, twigs and bark, fruits and flowers and debris. The litterfall inputs were about 20% lower at the eucalyptus site but this reduction was unevenly distributed among the litter categories. The reduction of the nitrogen and phosphorus inputs was larger (50%) than that of CPOM because of the low nutrient concentration of the CPOM at the eucalyptus site. Transported CPOM was also lower at the eucalyptus site. Although total CPOM inputs to the stream were reduced in the eucalyptus plantation, benthic storage of CPOM was 50% higher due to (1) high inputs of CPOM and low discharge during summer, (2) more twig and bark inputs, (3) eucalyptus leaves being retained more efficiently in the stream than deciduous leaves (4) a lower discharge, which may in part be attributable to eucalyptus-induced changes in the hydrological cycle. Increased retention balanced lower nitrogen and phosphorus content of CPOM, so benthic storage of nitrogen and phosphorus was similar at both sites. This work demonstrates that the timing, quality and quantity of inputs and benthic storage of CPOM in streams changes substantially because of the substitution of natural deciduous forest with eucalyptus plantation. Maintenance of buffer strips of natural vegetation may be the best way to protect ecological functioning of small, forested streams.     

Within-tree variation in cambial anatomy and xylem cell differentiation in Eucalyptus globulus

Summary Microscopic examination of the vascular cambium of Eucalyptus globulus, from four trees, at nine height levels, revealed a non-storied cambial structure with both ray and fusiform initials. Average fusiform initial length increased from 443 m at 70% of tree height from the base to a maximum of 484 m at 10% of tree height. At 2.5% of tree height average fusiform initial length sharply declined to 438 m. A strong positive correlation (r = 0.757, df = 7, P <0.05) existed between average fusiform initial length and average fibre length at each height, giving quantitative support to the notion that variation in fusiform initial length is an important mechanism influencing fibre length in hardwoods. However, 62% of the longitudinal variation in average fibre length was explained by the extent of elongation during differentiation, which increased linearly down the stem. At 70% of tree height, the average length of fibres was 1.84 times that of the fusiform initials, compared to 2.11 times at 2.5% of tree height. Concomitant with this longitudinal gradient of elongation was a gradient of increasing tangential area of the cambium occupied by cambial rays. This was accounted for by dimensional increases in the ray initial cells with decreasing height. Possible mechanisms that regulate the elongation of developing fibres and the proportion of cambial rays are discussed.     

Gall mite (Eriophyes laevis) infestation and leaf removal affect growth of leaf area in black alder (Alnus glutinosa) short shoots

Summary We measured the effects ofEriophyes laevis mite galls on the relative growth of short shoot leaf area ofAlnus glutinosa. A portion of leaves was artificially removed from a set of short shoots with both high and low gall density to cause local stress conditions. Nontreated high and low gall density short shoots were used as controls. The results show that the relative growth of leaf area measured for short shoots is negatively affected by high gall density. Artificial leaf removal, on the other hand, had positive effects on leaf area growth. Interestingly, the growth of leaf area did not differ for high gall density short shoots with leaf removal and noninfested short shoots with no leaf removal. This result may be caused by the combined, opposite effects of leaf removal and gall infestation.     

Variation of heartwood and sapwood in 18-year-old Eucalyptus globulus trees grown with different spacings

Heartwood and sapwood development was studied in 18-year-old Eucalyptus globulus trees from pulpwood plantations with different spacings (3 × 2, 3 × 3, 4 × 3, 4 × 4 and 4 × 5 m), on cross-sectional discs taken at breast height. The trees possessed a large proportion of heartwood, on average 60% of the wood cross-sectional surface. Spacing was a statistically significant source of variation of heartwood area, which ranged between 99 and 206 cm² for the closer (3 × 2) and wider (4 × 5) spacings, respectively. There was a positive and high statistical significant correlation between heartwood diameter and tree diameter (heartwood diameter = −0.272 + 0.616 dbh; r ² = 0.77; P < 0.001), and larger trees contained more heartwood regardless of spacing. Heartwood proportion in cross-section remained practically constant between spacings but increased with tree diameter class: 55.1, 62.2, 65.0 and 69.5% for diameter at breast height classes <15, 15–20, 20–25 and >25 cm, respectively. The sapwood width did not depend on tree diameter growth and remained practically constant at an average of 18 mm (range 15–21 mm), but sapwood area showed a good linear regression with tree diameter. Therefore, tree growth enhancement factors, such as wide spacings, will induce formation of larger heartwoods that can negatively impact raw-material quality for pulping. The increase in heartwood in relation with tree dimensions should therefore be taken into account when designing forest management guidelines.     

Genotype by environment interaction for growth of Eucalyptus globulus in Australia

This study assessed the genotype by environment (G × E) interaction for diameter growth in 15 Eucalyptus globulus progeny trials in Australia. Single-site analyses revealed significant subrace and family-within-subrace variance in all trials. Across-site subrace () and family () correlations were estimated by linear mixed model analyses of pairs of trials. Using a factor analytic structure for subrace and family random terms in a multi-environment mixed model analysis, best linear unbiased predictions of subrace effects were obtained for each trial. These were then averaged for each of four states (Victoria, Tasmania, South Australia and Western Australia) and across all sites. Statistically significant G × E interaction was detected, and weighted means across states for and were 0.73 and 0.76, respectively. Nevertheless, the three subraces from the Otway Ranges were both fast growing and relatively stable in their ranks over all sites. We evaluated the sensitivity of subraces to changing environmental conditions, on the basis of random coefficient models regressing subrace performance on selected trial climatic variables. The results suggested differential susceptibility of subraces to water, light and (to a less extent) temperature stresses during summer. Moreover, using multivariate techniques to visualize and interpret the across-site correlation structure for subrace effects, we could identify site clusters of reduced G × E interaction related to soil water availability and evaporative demand during summer. A revised site-type classification using these factors should allow a better capture of genetic gains from breeding and deployment.     

Growth dynamics of superficial roots in Portuguese plantations ofEucalyptus globulus Labill. studied with a mesh bag technique

Summary The variation in growth of the fine roots of blue gum (Eucalyptus globulus labill. ssp.globulus) in the 0–40 cm soil layer was studied from March 1982 to March 1983 at Quinta do Furaduoro, Óbidos, Portugal. A mesh bag method was used; bags of nylon net were inserted into a clay soil and a sandy soil and filled with root-free soil. They were resampled after 2, 4, 6 and 12 months in both places and, in a separate series in the sandy soil every second month throughout the year.The ingrowth of roots was high during the winter months but there was also a surprisingly high ingrowth during the spring-early summer period. There was also some root growth during the driest part of the yearviz. July–September.The amount of fine roots reached a maximum of about 260 g dw m^–2 after about 6 months in the sandy soil, whereas it took at least 12 months to reach the somewhat higher level of 450 g dw m^–2 in the clay soil. At that level the decomposition of dead roots was expected to equal the formation of new roots. Dead roots appeared after only 2 months. There was a higher proportion of dead roots in the clay soil than in the sandy soil, 35% as compared with 20% on an average, which indicates a slower decomposition or a higher mortality at equal decomposition rates in the clay than and in the sandy soil. The present data gives an indication of a minimum fine root production in mature Eucalyptus stands of at least 600 g dw m^–2 yr^–1.     

Genetic parameters for growth, wood density and pulp yield in Eucalyptus globulus

Genetic variation and co-variation among the key pulpwood selection traits for Eucalyptus globulus were estimated for a range of sites in Portugal, with the aim of improving genetic parameters used to predict breeding values and correlated response to selection. The trials comprised clonally replicated full-sib families (eight trials) and unrelated clones (17 trials), and exhibited varying levels of pedigree connectivity. The traits studied were stem diameter at breast height, Pilodyn penetration (an indirect measure of wood basic density) and near infrared reflectance predicted pulp yield. Univariate and multivariate linear mixed models were fitted within and across sites, and estimates of additive genetic, total genetic, environmental and phenotypic variances and covariances were obtained. All traits studied exhibited significant levels of additive genetic variation. The average estimated within-site narrow-sense heritability was 0.19 ± 0.03 for diameter and 0.29 ± 0.03 for Pilodyn penetration, and the pooled estimate for predicted pulp yield was 0.42 ± 0.14. When they could be tested, dominance and epistatic effects were generally not statistically significant, although broad-sense heritability estimates were slightly higher than narrow-sense heritability estimates. Averaged across trials, positive additive (0.64 ± 0.08), total genetic (0.58 ± 0.04), environmental (0.38 ± 0.03) and phenotypic (0.43 ± 0.02) correlation estimates were consistently obtained between diameter and Pilodyn penetration. This data argues for at least some form of pleiotropic relationship between these two traits and that selection for fast growth will adversely affect wood density in this population. Estimates of the across-site genetic correlations for diameter and Pilodyn penetration were high, indicating that the genotype by environment interaction is low across the range of sites tested. This result supports the use of single aggregated selection criteria for growth and wood density across planting environments in Portugal, as opposed to having to select for performance in different environments.     

Decomposition rate of Quercus rotundifolia leaves in an evergreen oak forest of the Duero Basin (Province of Zamora,Spain)

Experimental litter decomposition of Quercus rotundifolia leaves in an evergreen oak forest of the semiarid area of the Duero Basin has been carried out. The loss of weight of the decomposing oak leaves was 62% after 24 months. In general, it is physical factors (mainly precipitation) which limit the decomposition of the evergreen oak leaves. Different patterns of bioelement losses were observed, increasing the relative content of N and P, and decreasing the amounts of Mg, K and Na in the decomposing leaves.     

Effects of nickel on Frankia and its symbiosis with Alnus glutinosa (L.) Gaertn

The tolerance of nickel by Frankia in culture and in symbiosis with Alnus was determined. Yield of three Frankia strains was not affected significantly by 2.25 mM nickel when cultured in propionate medium containing hydolysed casein as nitrogen source. Yield of two strains in medium without combined nitrogen, and thus reliant on fixed nitrogen, was stimulated markedly by the same nickel concentration. Utilisation of nickel for synthesis of uptake hydrogenases is presumed to be the cause of enhanced nitrogenase activity.Although growth was reduced, treatment of 2-month-old seedlings with 0.025 mM nickel for 4 weeks did not affect nodulation significantly while nitrogenase activity was doubled. Nodulation and nitrogenase activity of seedlings receiving 0.075 mM nickel were inhibited markedly, while 0.5 mM nickel was lethal to all seedlings after 4 weeks of treatment. A few small, ineffective nodules were initiated early on some of the latter seedlings, suggesting that effects of nickel on host plant processes rather than Frankia are the primary cause of inhibition of nodulation. This interpretation is supported by the retention of substantial nitrogenase activity in 10-month-old plants 1 day after the treatment with 0.59 mM nickel, when the nickel content of roots and nodules was already maximal. No nitrogenase activity was detected after 3 days, by which time the leaves were almost completely necrotic. Over a 4 day period, most nickel was retained in the roots and nodules. Supplying histidine simultaneously at concentrations equal to, or in excess of, nickel prevented wilting and leaf necrosis, but did not increase translocation of nickel to the shoot.     

Decomposition of leaves of huisache (Acacia tortuoso) and mesquite (Prosopis spp) in soil of the central highlands of Mexico

In the central highlands of Mexico, mesquite (Prosopis spp) and huisache (Acacia tortuoso), N₂ fixing trees or shrubs, dominate the vegetation and are used in an alley cropping system to prevent erosion and restore soil fertility. We investigated how much the leaves of both trees contribute to dynamics of carbon (C) and nitrogen (N) in soil by adding leaves of both species to soil sampled under the canopy of mesquite and huisache, outside their canopy and from fields cultivated with maize at three different sites and monitoring microbial biomass C, production of carbon dioxide (CO₂), and dynamics of inorganic N (ammonium and nitrate) in an aerobic incubation. The soluble fraction and N content of the mesquite leaves were larger than in the huisache leaves, but lignin and polyphenol content were lower. Evolution of CO₂ increased 2.7-times when mesquite and 2.4-times when huisache leaves were added to soil. During all stages of decomposition and in all treatments, C mineralization of leaves from mesquite was greater than from huisache leaves. Mesquite leaves induced an increase in mineral N of 25.6 mg N kg^–1 soil after 56 days and those of huisache 9.8 mg N kg^–1. Twenty-six percent of N from mesquite leaves and 11% of huisache was mineralized, if no priming effect was considered. Nitrogen release from the leaves was greater when the soil organic matter content was lower. It was found that soil under the canopy of mesquite and huisache effectively accumulated organic material, micro-organisms and valuable nutrients. In an alley cropping system huisache might be a better choice than mesquite as huisache grows faster than mesquite and sheds its leaves twice a year while mesquite only once, although the amount of N mineralized was larger from mesquite leaves than from those of huisache.     

Effect of zinc supply on growth of three species of Eucalyptus seedlings and wheat

Summary Effects of zinc supply on shoot and root dry weight, root length, zinc concentrations and carbonic anhydrase activity were measured in 52 day old seedlings ofEucalyptus maculata, E. marginata, E. patens and wheat grown in a zinc deficient soil in the glasshouse.Symptoms of zinc deficiency in the eucalyptus and wheat appeared within 20 to 35 days. Eucalypt seedlings had short internodes and small necrotic leaves, reduced dry weight of shoots and roots, root length and zinc concentrations in young leaves; the measurable level of leaf carbonic anhydrase activity decreased to zero. Similar responses also occurred in wheat.The level of zinc fertilizer required for normal growth of Eucalyptus seedlings is therefore likely to be similar to that used for wheat and other agricultural crops.     

Movement patterns of an Australian chrysomelid beetle in a stand of two Eucalyptus host species

Summary We examined the movements of Chrysophtharta hectica, a eucalypt-feeding chrysomelid beetle in New South Wales, Australia, in relation to the beetle's sex, age and life-history, and to attributes of its Eucalyptus host plants. Beetle movements within the site were not influenced by beetle age or sex, but may be related to generation. Beetle distributions on the two host plant species, Eucalyptus stellulata and E. pauciflora, were generally clumped. Some of this clumping resulted from preference for E. stellulata over E. pauciflora. Clumping of beetles also occurred within host plant species; some plant individuals were consistently heavily used by beetles over the course of three years. We examined nutritional, spatial and biomass attributes of plants and found plant height and foliage production to be the best predictors of beetle numbers.     

Epidermal microstructures of the leaf,prophyll and nut in the himalayan species ofKobresia (Cyperaceae)

The epidermal structures of the foliage leaves, prophylls (perigynia) and nuts of 26 species and one variety ofKobresia (24 species and one variety from the Himalaya and two species from outside Himalaya) were studied by a scanning electron microscope (SEM) with an aim of making clear the relationship among these species. Most of the epidermal structures of the leaf and prophyll showed generic or familial relationships. The epidermal structures of the nuts were found useful for showing inter-specific relationships among morphologically closely related species though these structures were not found to be consistent with the infrageneric classification ofKobresia recognized by Clarke (1894, 1908) and Kükenthal (1909). On the basis of the epidermal structures of the nuts revealed by SEM the species ofKobresia could be put into four groups.     

Litterfall,leaf decomposition and litter colonization of Tessaria integrifolia (compositae) in the Parana river floodplain

Litterfall and leaf decomposition rates were measured in Choui Island, 45 km downstream from the confluence of the Paraná and Paraguay rivers. The material was collected biweekly from April 1985 through September 1986. Decomposition was measured in situ by the litter bag technique.Annual litterfall of Tessaria integrifolia gallery forest measured in the period April 1985 to March 1986 was 8.15 t ha^-1. Leaf litterfall was seasonal, i.e. significantly less leaf litter was shed during the high water phase than during the low water phase. The half life of the T. integrifolia litter over 38 days of decomposition was 20 days. At the beginning of the experiment, 15 and 38 days subsamples of remaining detritus were analyzed in order to determine changes in the nutrient content. After 38 days of incubation, the order of nutrient disappearance was Ca > K > N > Mg > Na > P.The number of invertebrates per g remaining litter of Tessaria integrifolia increased between incubations days 7 and 31. Collector-gatherers were more abundant after 38 days incubation; there were no shredders colonizing the leaf litter bags.     

Decomposition of Acer Rubrum Leaves at three depths in a eutrophic Ohio lake

Decomposition of terrestrial litterfall, that could enter a lake, was investigated at three depths within the lake in question using leaves of red maple (Acer rubrum). Sixty litter bags each containing 10 g dry weight of maple leaves were anchored at 1 m, 4 m and 12 m in East Twin Lake. At monthly intervals five bags were harvested at each level and dry weight changes noted. Results indicate little difference in decomposition between depths over the winter months. Beginning in May, however, significant differences were noted between 1 m and all other depths for the remainder of the experimental period. Maples lost 100% of weight at 1 m, 54% at 4 m, and 55% at 12 m over the experimental period. The rate of decomposition over the total experiment period was approximately .03 g da^–1 at 1 m and .015 g da^–1 at the 4 and 12 m depths. These results are discussed in terms of regulation of decomposition, position of decomposition and contribution to lake metabolism.     

Decomposition and colonisation by invertebrates of native and exotic leaf material in a small stream in New England (Australia)

In many parts of south-eastern Australia, native riparian vegetation has been cleared and exotic willows planted. In order to evaluate some of the possible effects of this practice, the decomposition and colonisation by invertebrates of the leaves of three native plant species along with those of willow were examined.Decomposition of leaves of the willow Salix babylonica L. and the indigenous macrophyte Myriophyllum propinquum A. Cunn. was much faster than for leaves of the indigenous trees Eucalyptus blakelyi Maiden and Casuarina cunninghamiana Miq. Both macroinvertebrates and current were found to have a significant influence upon decomposition. The pattern of preferential colonisation suggested that plant detritus represented a primary food source for invertebrates and not simply a refuge. Colonisation was found to be a function of the stage of decomposition, regardless of plant species. The lower temporal availability of willow leaves compared to the native evergreen tree leaves appears to be insufficient to enhance the production of the benthic macroinvertebrates.