Effects of nitrogen source and ectomycorrhizal association on growth and delta N-15 of two subtropical Eucalyptus species from contrasting ecosystems

Ectomycorrhizal (EM) associations facilitate plant nitrogen (N) acquisition, but the contribution of EM associations to tree N nutrition is difficult to ascertain in ecosystems. We studied the abilities of subtropical EM fungi and nutritionally contrasting Eucalyptus species, Eucalyptus grandis W. Hill ex Maiden and Eucalyptus racemosa Cav, to use N sources in axenic and soil cultures, and determined the effect of EM fungi on plant N use and plant N-15 natural abundance (delta N-15). As measured by seedling growth, both species showed little dependence on EM when growing in the N-rich minerotrophic soil from E. grandis rainforest habitat or in axenic culture with inorganic N sources. Both species were heavily dependent on EM associations when growing in the N-poor, organotrophic soil from the E. racemosa wallum habitat or in axenic culture with organic N sources. In axenic culture, EM associations enabled both species to use organic N when supplied with amide-, peptide- or protein-N. Grown axenically with glutamine- or protein-N, delta N-15 of almost all seedlings was lower than source N. The delta N-15 of all studied organisms was higher than the N source when grown on glutathione. This unexpected N-15 enrichment was perhaps due to preferential uptake of an N moiety more N-15-enriched than the bulk molecular average. Grown with ammonium-N, the delta N-15 of non-EM seedlings was mostly higher than that of source N. In contrast, the delta N-15 of EM seedlings was mostly lower than that of source N, except at the lowest ammonium concentration. Discrimination against N-15 was strongest when external ammonium concentration was high. We suggest that ammonium assimilation via EM fungi may be the cause of the often observed distinct foliar delta N-15 of EM and non-EM species, rather than use of different N sources by species with different root specialisations. In support of this notion, delta N-15 of soil and leaves in the rainforest were similar for E. grandis and co-occurring non-mycorrhizal Proteaceae. In contrast, in wallum forest, E. racemosa leaves and roots were strongly N-15-depleted relative to wallum soil and Proteaceae leaves. We conclude that foliar delta N-15 may be used in conjunction with other ecosystem information as a rapid indicator of plant dependency on EM associations for N acquisition.

Physiological responses to water stress in Eucalyptus cloeziana and E-argophloia seedlings

Effects of water stress duration and intensity on gas exchange and leaf water potential were investigated in 7-month-old seedlings of a humid coastal provenance (Gympie) and a dry inland (Hungry Hills) provenance of E. cloeziana F. Muell. and in a dry inland (Chinchilla) provenance of E. argophloia Blakely supplied with 100% (T-100), 70% (T-70), 50% (T-50) of their water requirements, or were watered only after they were wilted at dawn (T-0). Seedlings of E. argophloia had the highest midday net photosynthetic rate (A), stomata] conductance (g(s)), stomatal density and predawn leaf water potential (Psi(pd)) in all treatments. The E. cloeziana provenances did not differ in these attributes. The T-70 and T-50 treatments caused reductions in A of 30% in E. argophloia, and 55% in the E. cloeziana provenances. Under the T-0 treatment, E. argophloia maintained higher rates of gas exchange at all levels of water stress than E. cloeziana provenances. The estimates of Psi(pd) and midday water potential (Psi(md)) at which plants remained wilted overnight were respectively: -2.7 and -4.1 MPa for E. cloeziana (humid), -2.8 and -4.0 MPa for E. cloeziana (dry) and, -3.7 and -4.9 MPa for E. argophloia. Following stress relief, both A and g(s) recovered more quickly in E. argophloia and in the dry provenance of E. cloeziana than in the humid provenance. We conclude that E. argophloia is more drought tolerant and has a potential for cultivation in the humid and semi humid climates, whilst E. cloeziana has greater potential in the humid subtropical climates.

Selection of species and provenances for low-rainfall areas: physiological responses of Eucalyptus cloeziana and Eucalyptus argophloia to seasonal conditions in subtropical Queensland

Responses of stomatal conductance (g(s)) and net photosynthesis (A) to changes in soil water availability, photosynthetic photon flux density (Q), air temperature (1) and leaf-to-air vapour pressure deficit (D) were investigated in 4-year-old trees of a dry inland provenance of Eucalyptus argophloia Blakely, and two dry inland provenances (Coominglah and Hungry Hills) and a humid coastal provenance (Wolvi) of Eucalyptus cloeziana F. Muell. between April 2001 and April 2002 in southeast Queensland, Australia. There were minimal differences in A, g, and water relations variables among the coastal and inland provenances of E. cloeziana but large differences between E. argophloia and E. cloeziana. E. argophloia and to a lesser extent the Hungry Hills (inland) provenance of E. cloeziana maintained relatively higher pre-dawn water potential (psi(pd)) during the dry season suggesting possible access to water at depth. Simple phenomenological models of stomatal conductance as a function of Q, T and D explained 60% of variation in gs in E. cloeziana and more than 75% in E. argophloia, when seasonal effect was incorporated in the model. A Ball-Berry model for net photosynthesis explained between 70 and 80% of observed variation in A in both species. These results have implications in matching the dry and humid provenances of E. cloeziana and E. argophloia to suitable sites in subtropical environments. (C) 2004 Elsevier B.V. All rights reserved.

Reciprocal N ((NH4+)-N-15 or (NO3-)-N-15) transfer between nonN(2)-fixing Eucalyptus maculata and N-2-fixing Casuarina cunninghamiana linked by the ectomycorrhizal fungus Pisolithus sp.

Two-way N transfers mediated by Pisolithus sp. were examined by excluding root contact and supplying (NH4+)-N-15 or (NO3-)-N-15 to 6-month-old Eucalyptus maculata or Casuarina cunninghamiana grown in two-chambered-pots separated by 37 m screens. Mycorrhizal colonization was 35% in Eucalyptus and 66% in Casuarina (c. 29% N-2-fixation). Using an environmental scanning electron microscope, living hyphae were observed to interconnect Eucalyptus and Casuarina. Biomass and N accumulation was greatest in nodulated mycorrhizal Casuarina/mycorrhizal Eucalyptus pairs, less in nonnodulated mycorrhizal Casuarina/mycorrhizal Eucalyptus pairs, and least in nonnodulated nonmycorrhizal Casuarina/nonmycorrhizal Eucalyptus pairs. In nonnodulated mycorrhizal pairs, N transfers to Eucalyptus or to Casuarina were similar (2.4-4.1 mg per plant in either direction) and were 2.6-4.0 times greater than in nonnodulated nonmycorrhizal pairs. In nodulated mycorrhizal pairs, N transfers were greater to Eucalyptus (5-7 times) and to Casuarina (12-18 times) than in nonnodulated mycorrhizal pairs. Net transfer to Eucalyptus or to Casuarina was low in both nonnodulated nonmycorrhizal (< 0.7 mg per plant) and nonnodulated mycorrhizal pairs (< 1.1 mg per plant). In nodulated mycorrhizal pairs, net transfer to Casuarina was 26.0 mg per plant. The amount and direction of two-way mycorrhiza-mediated N transfer was increased by the presence of Pisolithus sp. and Frankia, resulting in a net N transfer from low-N-demanding Eucalyptus to high-N-demanding Casuarina.

Phenolic acids and abscisic acid in Australian Eucalyptus honeys and their potential for floral authentication

Seven phenolic acids related to the botanical origins of nine monofloral Eucalyptus honeys from Australia, along with two abscisic isomers, have been analyzed. The mean content of total phenolic acids ranges from 2.14 mg/100 g honey of black box (Eucalyptus largiflorens) honey to 10.3 mg/100 g honey of bloodwood (Eucalyptus intermedia) honey, confirming an early finding that species-specific differences of phytochemical compositions occur quantitatively among these Eucalyptus honeys. A common profile of phenolic acids, comprising gallic, chlorogenic, coumaric and caffeic acids, can be found in all the Eucalyptus honeys, which could be floral markers for Australian Eucalyptus honeys. Thus, the analysis of phenolic acids could also be used as an objective method for the authentication of botanical origin of Eucalyptus honeys. Moreover, all the honey samples analyzed in this study contain gallic acid as the main phenolic acid, except for stringybox (Eucalyptus globoidia) honey which has ellagic acid as the main phenolic acid. This result indicates that the species-specific differences can also be found in the honey profiles of phenolic acids. Further-more, the analysis of abscisic acid in honey shows that the content of abscisic acid varies from 0.55 mg/100 g honey of black box honey to 4.68 mg/ 100 g honey of bloodwood honey, corresponding to the contents of phenolic acids measured in these honeys. These results have further revealed that the HPLC analysis of honey phytochemical constituents could be used individually and/or jointly for the authentication of the botanical origins of Australian Eucalyptus honeys. (C) 2003 Elsevier Ltd. All rights reserved.

Growth performance and management of a mixed rainforest tree plantation

Monoculture plantations of Pinus, Eucalyptus and Acacia have been established oil rainforest lands throughout the world. However, this type of reforestation generally supplies low quality timber and contributes to landscape simplification. Alternatives to exotic monoculture plantations are now beginning to gain momentum with farmers and landholders attempting to establish a variety of rainforest trees in small plantations. When compared to the well studied commercial species, knowledge concerning the growth and management of many of these rainforest species is in its infancy. To help expand this limited knowledge base an experimental plantation of 16 rainforest tree species in a randomised design was established near Mt. Mee, in south-eastern Queensland, Australia. Changes in growth, form (based on stem straightness, branch size and branchiness), crown diameters and leaf area of each species were examined over 5 years. Patterns of height growth were also measured monthly for 31 months. Species in this trial could be separated into three groups based on their overall growth after 5 years and their growth patterns. Early successional status, low timber density, high maximum photosynthetic rates and large total leaf areas were generally correlated to rapid height growth. Several species (including Araucaria cunninghamii, Elaeocarpus grandis, Flindersia brayleyana, Grevillea robusta and Khaya nyasica) had above average form and growth, while all species in the trial had considerable potential to have increased productivity through tree selection. As canopy closure occurred at the site between years four and five, growth increments declined. To reduce stand competition a number of different thinning techniques could be employed. However, simple geometric or productivity based thinnings appear to be inappropriate management techniques for this mixed species stand as they would either remove many of the best performing trees or nearly half the species in the trial. Alternatively, a form based thinning would maintain the site's diversity, increase the average form of the plantation and provide some productivity benefits.

Nodulated N-2-fixing Casuarina cunninghamiana is the sink for net N transfer from non-N-2-fixing Eucalyptus maculata via an ectomycorrhizal fungus Pisolithus sp using (NH4+)-N-15 or (NO3-)-N-15 supplied as ammonium nitrate

To determine the effects of nitrogen source on rates of net N transfer between plants connected by a common mycorrhizal network, we measured transfer of N supplied as (NH4NO3)-N-15-N-14 or (NH4NO3)-N-14-N-15 in three Casuarina/Eucalyptus treatments interconnected by a Pisolithus sp. The treatments were nonnodulated nonmycorrhizal/nonmycorrhizal; nonnodulated mycorrhizal/mycorrhizal; and nodulated mycorrhizal/mycorrhizal. Mycorrhization was 67% in Eucalyptus and 36% in Casuarina. N-2 fixation supplied 38% of the N in Casuarina. Biomass, N and N-15 contents were lowest in nonmycorrhizal plants and greatest in plants in the nodulated/mycorrhizal treatment. Nitrogen transfer was enhanced by mycorrhization and by nodulation, and was greater when N was supplied as (NH4+)-N-15 than (NO3-)-N-15. Nitrogen transfer rates were lowest in the nonmycorrhizal treatment for either N-15 source, and greatest in the nodulated, mycorrhizal treatment. Transfer was greater to Casuarina than to Eucalyptus and where ammonium rather than nitrate was the N source. Irrespective of N-15 source and of whether Casuarina or Eucalyptus was the N sink, net N transfer was low and was similar in both nonnodulated treatments. However, when Casuarina was the N sink in the nodulated, mycorrhizal treatment, net N transfer was much greater with (NH4+)-N-15 than with (NO3-)-N-15. High N demand by Casuarina resulted in greater net N transfer from the less N-demanding Eucalyptus. Net transfer of N from a non-N-2-fixing to an N-2-fixing plant may reflect the very high N demand of N-2-fixing species.

Modelling predicts positive and negative interactions between three Australian tropical tree species in monoculture and binary mixture

Computer modelling promises to. be an important tool for analysing and predicting interactions between trees within mixed species forest plantations. This study explored the use of an individual-based mechanistic model as a predictive tool for designing mixed species plantations of Australian tropical trees. The 'spatially explicit individually based-forest simulator' (SeXI-FS) modelling system was used to describe the spatial interaction of individual tree crowns within a binary mixed-species experiment. The three-dimensional model was developed and verified with field data from three forest tree species grown in tropical Australia. The model predicted the interactions within monocultures and binary mixtures of Flindersia brayleyana, Eucalyptus pellita and Elaeocarpus grandis, accounting for an average of 42% of the growth variation exhibited by species in different treatments. The model requires only structural dimensions and shade tolerance as species parameters. By modelling interactions in existing tree mixtures, the model predicted both increases and reductions in the growth of mixtures (up to +/- 50% of stem volume at 7 years) compared to monocultures. This modelling approach may be useful for designing mixed tree plantations. (c) 2006 Published by Elsevier B.V.

Effect of NaCl on emergence and growth of a range of provenances of Eucalyptus citriodora, Eucalyptus populnea, Eucalyptus camaldulensis and Acacia salicina

High salt levels in mine spoils have been identified as one of the major chemical limitations to plant establishment after coal mining in central Queensland. Soil solution extracts from spoils indicated that EC levels of up to 26 dS/m could be encountered. Glasshouse trials examined the emergence and growth of Eucalyptus citriodora, Eucalyptus camaldulensis and Eucalyptus populnea provenances and Acacia salicina subjected to such EC levels. Relatively low levels of salt (100 mM NaCl, or 11 dS/m) with respect to the levels encountered on mine spoils, were enough to substantially reduce the rate and percentage emergence of all eucalypt provenances. A. salicina was found to be superior to the eucalypts in its ability to emerge and survive under saline conditions. It was the only species to have seedlings emerge and survive at 200 mM NaCl (20 dS/m), and the effect of salt on decreasing seedling dry weight was less pronounced for A. salicina than for any of the eucalypts. Established plants survived the range of salt treatments far better than emerging seedlings, with survival of established plants being reduced only at 300 and 400 mM NaCl (28 and 36 dS/m, respectively). A. salicina performed significantly better at 300 and 400 mM NaCl than most of the eucalypts studied. (c) 2006 Elsevier B.V. All rights reserved.