Phosphorus accumulation by lignotubers of jarrah (Eucalyptus marginata Donn ex Sm.) seedlings grown in a range of soils

Summary Jarrah seedlings were grown in six virgin Western Australian soils for up to 27 months. Lignotubers were produced in all soils and formed 10–16% of plant dry weight. The phosphorus concentration in the lignotuber (250–800 g g^–1) was nearly twice that in the stem and roots. The lignotuber contained 10–30% of total plant phosphorus and like the leaves was a sink for phosphorus. In one lateritic soil the phosphorus concentrations of lignotuber and stem barks were similar. However, in the same plants the concentration of phosphorus in the lignotuber wood was five times the phosphorus concentration in stem wood. Hence both lignotuber bark and sap wood in young jarrah seedlings are storage sites for phosphorus. X-ray probe analysis showed that wood phosphorus was associated with the ray parenchyma. Unlike phosphorus, nitrogen did not accumulate in the lignotuber and the concentrations of nitrogen were similar for roots, lignotubers and stems.     

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.     

Tracing water uptake by jarrah (Eucalyptus marginata) trees using natural abundances of deuterium

 Seasonal change in the δ²H content of water from twig sap, soil, rainfall and groundwater were measured to determine the water sources accessed by jarrah (Eucalyptus marginata) trees at three sites in Western Australia with differing soils and depths to water table. During winter and spring the main contributor to the water uptake of the trees was stored water in the surface layers of the soil replenished by predominantly winter rainfall. With the onset of summer drought jarrah became more reliant on water from deeper down the profile. There was no clear evidence that jarrah could tap water from groundwater more than 14 m deep in deep sands. Defining the source of water for trees in deep lateritic soils using stable isotopes is hampered by the uniform deuterium profiles down most of the unsaturated zone and into the groundwater. There was a limited response in the δ²H values of sapwater in twigs to changes in the δ²H of the upper layers of the deep sand following input of rainfall in autumn. The damped response was related to the small variation in the δ²H composition of rainfall in most events during the year and the mixing in the tree of water extracted from different locations in the soil profile. Received: 21 August 1995 / Accepted: 3 December 1995     

Processing of leaf litter in a northern jarrah forest stream,Western Australia: I. Seasonal differences

Seasonal differences in the processing of jarrah (Eucalyptus marginata) leaves were examined in a small forest stream of the Darling Range, Western Australia. Processing rates in both summer and winter were very low, placing this species in the slow category (k < 0.005) of Petersen & Cummins (1974).Macroinvertebrates did not contribute to the processing of jarrah leaves during summer and autumn, coincident with the period of peak litter fall. The low quality of detritus entering the streams, and possibly some adverse interactions with the physical and chemical environment, inhibits processing during the summer months. As a consequence, a significant proportion of the coarse-particle detritus entering these streams may be exported unprocessed at the onset of high winter flows.In contrast with previous studies, jarrah leaves were processed at a faster rate during winter than summer. This seasonal difference can, in part, be attributed to the increased density and proportion of shredders at this time. Despite this, the processing of jarrah leaves during winter was very slow and must reflect the poor quality of this detritus as food for invertebrate consumers.     

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.     

Effects of root-zone solutes on Eucalyptus camaldulensis and Eucalyptus bicostata seedlings: Responses to Na+, Mg2+ and Cl-

Specific-ion effects in salt-treated eucalypts were examined with two species known to differ in salt tolerance viz. E. camaldulensis (more tolerant) and E. bicostata (less tolerant). Sand-cultured plants were irrigated with different nutrient solutions designed to impose either osmotic stress (concentrated macronutrients with balanced cations and anions) or specific ion stress from either NaCl or MgCl₂, or from nutrient solutions rich in particular ions viz. Na⁺, Mg²⁺ and Cl^- (balancing counter ions were provided in all cases). Half-strength Hoagland nutrient solution served as control. All treatments were applied at osmotic pressures of approximately 0.52 MPa by appropriate concentrations of each solution. In general, salt-induced growth reductions were greater for E. camaldulensis than for E. bicostata, although E. camaldulensis showed strongest exclusion of Na⁺, Mg²⁺ and Cl^- from shoots. Application of NaCl and concentrated macronutrients resulted in similar growth reductions. E. bicostata seedlings exposed to high Cl^- concentrations in the presence of Mg²⁺ and concentrated cations suffered significantly more shoot and root reduction than those exposed to other salts. Treatment with solution rich in Cl^- resulted in extensive leaf damage, which suggested that Cl^- may have exerted a specific effect. No specific Na⁺ effect was observed for either species, even though shoot Na⁺ concentrations were considerably higher for E. bicostata than for E. camaldulensis. Root growth was considerably less for plants treated with Mg²⁺ salts and this effect was associated with low root Ca²⁺ concentrations.     

Boron requirements for Eucalyptus globulus seedlings

Although boron (B) deficiency limits the productivity of eucalypts in plantations in many parts of the world, the concentrations of foliar B used in the diagnosis of B deficiency vary greatly among studies. There has been a lack of reliable diagnosis standards for B deficiency in Eucalyptus species. Therefore, the present study investigated the relationship between internal and external B concentrations and growth of Eucalyptus globulus, the main commercial temperate eucalypt species. Seedlings were grown in a B-buffered solution culture (Amberlite IRA 743) from 0.03 to 8.35 M B. Boron deficiency symptoms appeared at day 5 in the nutrient solution containing less than 0.27 M B. The external critical B concentrations, estimated for the growth of shoots and roots, were 1.08 and 0.99 M B, respectively. The internal critical B concentration range in the youngest fully expanded leaf (YFEL) for shoot growth was 12–16 mg B kg^–1 dry weight. The internal critical B concentrations estimated in the present study have been successfully used in the diagnosis of B deficiency in E. globulus trees up to three years of age in south-east Asia.     

Unmated queens in the primitively eusocial waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Summary In the primitively eusocial tropical waspRopalidia marginata, five out of eleven colonies studied had an unmated female as their queen. In two colonies this was the case despite the presence of another mated individual in the colony. We found no detectable differences between colonies with unmated queens and those with mated queens. We argue that in species such asR. marginata, where intracolony relatedness is expected to be low and where sociality is likely to be maintained because several individuals have opportunities for direct reproduction in the future, individual selection is likely to override the good of the colony and lead to such phenomena as that of unmated queens.     

Queen succession in the primitively eusocial tropical waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Females of the primitively eusocial wasp Ropalidia marginata can be classified into three behavioral groups: Sitters, Fighters, and Foragers. It has been speculated that both Sitters and Fighters may be hopeful queens and that the Foragers may have little or no opportunities for direct reproduction. Here we show that in 9 of 12 queen-removal experiments where such a behavioral differentiation could be discerned, the individual that became a queen (the potential queen) was a Sitter in 6 cases, a Fighter in 2 cases, and a Forager in only 1 case. Although potential queens spent significantly more time absent from the nest and showed significantly higher rates of dominance behavior compared to the mean values for nonqueens in their colonies, they were intermediate with respect to all behaviors and age when compared to the range of values for nonqueens in their colonies. Potential queens were not necessarily the highest-ranking individuals among the nonqueens. The pattern of queen succession in this species appears to be quite different from the temperate pattern, where an old and active forager of high dominance rank is the potential queen. Although somewhat similar to the tropical pattern of a relatively younger female that has performed relatively little foraging being the potential queen, it is perhaps more accurate to describe the potential queens of R. marginata as unspecialized intermediates.     

Diagnosis of zinc deficiency in seedlings of a tropical eucalypt (Eucalyptus urophylla S. T. Blake)

A glasshouse experiment was undertaken to establish the internal Zn requirement for shoot growth of Eucalyptus urophylla, a fast-growing commercial plantation species widely planted in tropical regions of the world. A Zn-deficient sand was supplemented with ten rates of Zn and seedlings were harvested after three months. In Zn-deficient plants the new growth was dwarfed with small, necrotic leaves and short internodes. Foliar Zn concentrations declined markedly with leaf age in both Zn-deficient and Zn-adequate plants. The critical Zn concentration for the diagnosis of Zn deficiency also fell with leaf age. Zinc concentrations in the youngest fully expanded leaf ranged from 8–11 g Zn g^–1 dry weight in plants with severe symptoms to 30–37 g Zn g^–1 dry weight in non-deficient plants. The critical Zn concentration for the diagnosis of Zn deficiency at 90% of maximum shoot growth in the same leaf was 21 g Zn g^–1 dry weight. This value is nearly twice that reported for several other species of eucalypts and may indicate a higher internal demand for Zn in tropical than in temperate eucalypts.     

Ecto- and arbuscular mycorrhizal symbiosis can induce tolerance to toxic pulses of phosphorus in jarrah (Eucalyptus marginata) seedlings

In common with many plants native to low P soils, jarrah (Eucalyptus marginata) develops toxicity symptoms upon exposure to elevated phosphorus (P). Jarrah plants can establish arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) associations, along with a non-colonizing symbiosis described recently. AM colonization is known to influence the pattern of expression of genes required for P uptake of host plants and our aim was to investigate this phenomenon in relation to P sensitivity. Therefore, we examined the effect on hosts of the presence of AM and ECM fungi in combination with toxic pulses of P and assessed possible correlations between the induced tolerance and the shoot P concentration. The P transport dynamics of AM (Rhizophagus irregularis and Scutellospora calospora), ECM (Scleroderma sp.), non-colonizing symbiosis (Austroboletus occidentalis), dual mycorrhizal (R. irregularis and Scleroderma sp.), and non-mycorrhizal (NM) seedlings were monitored following two pulses of P. The ECM and A. occidentalis associations significantly enhanced the shoot P content of jarrah plants growing under P-deficient conditions. In addition, S. calospora, A. occidentalis, and Scleroderma sp. all stimulated plant growth significantly. All inoculated plants had significantly lower phytotoxicity symptoms compared to NM controls 7 days after addition of an elevated P dose (30 mg P kg^?1 soil). Following exposure to toxicity-inducing levels of P, the shoot P concentration was significantly lower in R. irregularis-inoculated and dually inoculated plants compared to NM controls. Although all inoculated plants had reduced toxicity symptoms and there was a positive linear relationship between rank and shoot P concentration, the protective effect was not necessarily explained by the type of fungal association or the extent of mycorrhizal colonization.     

Effects of aluminium on growth and cation uptake in seedlings of Eucalyptus mannifera and Pinus radiata

This experiment was designed to examine the effects of aluminium (Al) on the growth of Pinus radiata (D. Don) and Eucalyptus mannifera subsp. mannifera (Mudie) seedlings in culture solutions in a glasshouse to help explain the failure of radiata pine trees on some acid, low fertility soils in Australia on which the native eucalypts flourish. Aluminium (Al) in culture solution increased the growth of roots and shoots of seedlings of both species but while growth of the eucalypt continued to increase with increases in Al to 2.222 μM, growth of the pine was largest at 370 μM Al. In addition to total root length, specific root length (length per unit dry weight), a measure of fineness of the root, increased in the eucalypt seedlings as the substrate Al increased. Growth of the shoots and roots of the pine in the absence of any added Al was extremely poor suggesting that Al, in low concentrations, may be an essential element or ameliorate some other factors in solution culture at low pH. Root and shoot concentrations of K increased with increasing Al, whilst Ca and Mg Concentrations decreased and Mn concentrations were unaffected in both species. Tissue Ca and Mg concentrations were 2 to 3 times higher in the eucalypt seedlings than the pine at all levels of added Al due to greater uptake of these elements by the eucalypt. In contrast, at the highest concentration of Al in the medium, shoot Al concentrations were lower in the cucalypt than in the pine due to a greater proportion of Al being retained in the eucalypt roots. These differences between the seedlings in terms of root growth and tissue cation concentrations may help explain the ability of eucalypt species to maintain vigorous growth on acid soils high in Al and low in Ca and P, where growth of the pines failed.     

Emergence of Eucalyptus marginata (jarrah) from seed in Mediterranean-climate forest in response to overstorey,site, seedbed and seed harvesting

Abstract The effects of overstorey, type of site, seedbed, seeding date and seed harvesting by vertebrates and invertebrates on the emergence of Eucalyptus marginata (jarrah) seedlings were studied experimentally in the northern jarrah forest of Western Australia. Seed harvesting by small vertebrates substantially reduced emergence, whereas invertebrates only slightly reduced emergence. Ninety per cent of seed was removed within 1 day when it was on the soil surface and in the open, whereas seed removal was insignificant when the seed was covered by soil. Vertebrates harvested seed more effectively when the understorey and litter were removed and the seed was more visible on the soil surface. Seed harvesting by small vertebrates had less impact on a site where there had been bauxite mining, possibly because the surrounding area was being revegetated with large numbers of seed and the area afforded poor cover for vertebrates. Emergence was less where the overstorey was removed compared to where it was retained, and on the rehabilitated bauxite mined site compared to the forest sites. There was no difference in emergence between the low and high quality forest sites and so the emergence phase could not fully explain the variation in abundance off. marginata seedlings on different quality sites. Post emergence events are likely to be important in explaining this difference between low and high quality sites.     

Copper nutrition of Eucalyptus maculata Hook. seedlings: Requirements for growth,distribution of copper and the diagnosis of copper deficiency

A glasshouse experiment was conducted to define the response of Eucalyptus maculata seedlings to the addition of nine rates of copper (Cu) to a Cu-deficient sand. Plants were harvested 128 days from sowing. Symptoms of Cu deficiency included marginal necrosis in young fully expanded leaves (YFEL), deformed leaf margins, death of lateral shoots, bleeding at nodes on the main stem and reduced lignification of xylem fibres and vessels. Plant height and the number of nodes on the main stem were unaffected. In plants supplied with 0 Cu, whole top and root fresh weights were depressed by 27% and 32% respectively. The external Cu requirement for maximum growth of E. maculata seedlings was similar to that for wheat grown in the same soil. In Cu-adequate plants, leaf Cu concentrations decreased with distance from the shoot apex. Cu levels in stems varied little with position and were similar to the YFEL. Cu concentrations in leaves and stems were depressed in Cu-deficient plants to <1.0 g g^–1 dry weight (d.w.) (roots: 1.5 g g^–1 d.w.). The external Cu supply did not greatly alter the distribution of Cu within the plant. Young leaves at the shoot tip are recommended for diagnosis of Cu deficiency: critical values for shoot d.w. were about 1.5 g Cu g^–1 d.w. Lignification of wood was suppressed where Cu concentrations fell below 1.5 g g^–1 d.w.: the Bussler test for lignification would thus be a valuable indicator of Cu deficiency.     

Decomposition of Eucalyptus globulus leaves and three native leaf species (Alnus glutinosa, Castanea sativa and Quercus faginea) in a Portuguese low order stream

Leaf decomposition of the exotic evergreen Eucalyptus globulus (eucalyptus), and three native deciduous tree species, Alnus glutinosa (alder), Castanea sativa (chestnut) and Quercus faginea (oak), was compared in a second order stream in Central Portugal. Changes in dry weight, nitrogen and polyphenolic compounds and microbial colonization were periodically assessed for three months.Negative exponential curves fit the leaf weight loss with time for all leaf species. Mass loss rate was in the order alder (K = 0.0161) > chestnut (K = 0.0079) > eucalyptus (K = 0.0068) > oak (K = 0.0037). Microbial colonization followed the same pattern as breakdown rates. Evidence of fungal colonization was observed in alder after 3 days in the stream, whereas it took 21 days in oak leaves to have fungal colonization. Fungal diversity was leaf species-dependent and increased with time. In all cases, percent nitrogen per unit leaf weight increased, at least, at the initial stages of decay while soluble polyphenolics (expressed as percentage per unit leaf weight) decreased rapidly in the first month of leaves immersion.Intrinsic factors such as nitrogen and polyphenolic content may explain differences in leaf decomposition. The possible incorporation of eucalyptus litter into secondary production in a reasonable time span is suggested, although community balance and structure might be affected by differences in allochthonous patterns determined by eucalyptus monocultures.     

Factors affecting the acceptance of alien conspecifics on nests of the primitively eusocial wasp,Ropalidia marginata (Hymenoptera: Vespidae)

In the primitively eusocial wasp, Ropalidia marginata,individual females are known to drift from one newly founded nest to another. In the laboratory, young (<6- to 8-day-old) alien wasps are accepted onto unrelated colonies, while older (>6- to 8-day-old) wasps are not. Here we have investigated the factors that could influence the acceptance of foreign conspecifics onto unrelated nests. Individually marked wasps of different ages, isolated immediately after eclosion from the natal nest and from each other, were introduced onto unrelated recipient nests. Considered separately, both age and ovarian condition seemed to influence the probability of acceptance as well as the levels of aggression and tolerance received by the introduced wasps. However, partial correlation analysis and multiple regression analysis indicated that only age had a direct influence and that the ovarian condition acts only through age, a variable with which it is highly correlated. The observed acceptance of young aliens and rejection of old aliens are less likely to be due to the perception of older wasps as a reproductive threat rather than some age-related factor, other than ovarian condition, for example, the relative ease with which younger wasps can be molded into desired roles.     

Study of endophytic fungal community from different parts of <Emphasis Type="Italic">Aegle marmelos</Emphasis> Correae (Rutaceae) from Varanasi (India)

Endophytic fungi were isolated from healthy, living, and symptomless tissues of inner bark, leaf, and roots of Aegle marmelos, a well-known medicinal plant, growing in different parts of India including Varanasi. A total of 79 isolates of endophytic fungi were isolated, representing 21 genera, adopting a standard isolation protocol. Members of the deuteromycotina were more prevalent than ascomycotina and others. The result was quite encouraging in terms of maximum isolates recovery from hyphomycetes (78.5%) followed by ascomycetes (8.9%) and coelomycetes (7.6%) respectively, which corroborates previous studies in same area. However, 5.1% isolates remained unidentified and were classified under Mycelia Sterilia. No isolate was obtained from either basidiomycotina or from zygomycotina. Fusarium spp. had maximum colonization frequency (8.00%) in this plant. The other dominant endophytic genera were Aspergillus spp., Alternaria sp., Drechslera sp., Rhizoctonia sp., Curvularia sp., Nigrospora sp., and Stenella sp. Only two ascomycetous members Chaetomium globosum and Emericella sp. (perfect state of Aspergillus sp.) were obtained from the bark sample. These results indicated that distribution of endophytic fungi within the A. marmelos is not even. Bark harbors more endophytic fungi than leaf and root.     

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.     

Growth and nutrition of birch seedlings in relation to potassium supply rate

Summary Growth of hydroponically cultivated birch seedlings (Betula pendula Roth.) at sub- and supra-optimum potassium supply rates was investigated. Potassium was supplied either as a relative addition rate (r _k = 5, 10, 15 and 20% increase day^-1) or as fixed concentrations (0.2, 3, 6, 12 and 15 mM) in the culture solution. After an acclimation period the growth rate of the seedlings in the suboptimum treatments reached values close to the treatment variable, the relative rate of K-addition. Deficiency symptoms, in the form of chlorosis and necroses along the leaf margins, developed initially in all suboptimum treatments, but very few new symptoms appeared once the seedlings had reached the phase of steady-state nutrition and growth. At supra-optimum K-supply levels, i.e. at 0.2–15 mM K in the culture solution, no symptoms of deficiency or toxicity developed, and the relative growth rate of the seedlings remained maximum. The relative growth rate of the seedlings was linearly related to the plant K-status for K contents ranging from 0.2 to 1.0% of dry weight (DW). At higher internal K-concentrations, 1.0–3.0% DW, no further increase in relative growth rate was achieved. A shortage of K resulted in a decrease in the net assimilation rate. This effect was counterbalanced by the absence of shift in he leaf weight ratio as well as by the production of relatively thin leaves. The fraction of dry matter allocated to roots decreased in K-limited plants, as did the leaf contents of soluble carbohydrates and starch.     

The effect of soil desiccation on the nutrient status of Eucalyptus regnans F. Muell seedlings

The poor growth of young Eucalyptus regnans seedlings in undried soil from the mature forest of E. regnans can be overcome by previously air-drying the soil or by adding sufficient amounts of complete soluble fertilizer or equivalent concentrations of P (as NaH₂PO₄) and N (as NaNO₃). A factorial pot experiment in which phosphate and nitrate were added to undried soil indicated that P was the primary deficiency for young seedlings and that response to N did not occur until this lack was satisfied. In dried soil, seedlings also responded to additions of complete fertilizer but most of this effect was due to N rather than P. Field trials in the mature forest also indicated greater growth in dried soil than undried soil and confirmed a response of young seedlings to superphosphate. In pot experiments, the concentration of P and N per g plant dry weight after four months was relatively constant irrespective of the final size of the plant. Seedlings in dried soil extracted up to 15 times more P than did those grown in undried soil. In general, chemical analysis of soil indicated more extractable P and N from dried soil although this was not always consistently so. Soil desiccation resulted in an increase in soil surface area due to the fragmentation of larger peds and to an increase in the number of microfractures which remained in the soil crumbs after rewetting. Mycorrhiza are likely to be important since the differentiation of the growth response of seedlings in dried and undried soil, which occurred at 5–6 weeks, corresponded with the establishment of full ectomycorrhizal development (80% root tips). The factors concerned with the increase in fertility after air-drying are discussed.Abbreviations GR Growth Ratio