ALBERT

Notes: The foliar natural abundance of 15N was analysed to compare the potential nitrogen sources of vascular rainforest epiphytes and associated soil-rooted trees. Leaves of epiphytes collected from six rainforest communities in Brazil, Australia and the Solomon Islands were depleted in 15N relative to the trees at each site. Epiphyte δ15N was as low as -6.4%o, while trees were generally enriched in 15N (0.7 to 3.5%o). These results indicate either that epiphytes use nitrogen sources depleted in 15N or that discrimination against 15N is an intrinsic function of epiphyte physiology. At three sites, epiphytes could be grouped into those having both low δ15N and low leaf-nitrogen content and those possessing both high δ15N and high leaf-nitrogen content. The second group had δ15N values in the range sometimes attributable to N2 fixation (-2 to 0%o). There was no correlation between growth form and δ15N. It is concluded that epiphytes may utilize 15N-depleted nitrogen from atmospheric deposition and N2 fixation.

Notes: Abstract Growth and gas exchange measurements are used in conjunction with a carbon balance model to describe the millet (Pennisetum typhoides)–witchweed (Striga hermonthica) host—parasite association. Striga hermonthica reduces the growth of millet by 28% and radically alters the architecture of infected plants. Whilst grain yield and stem dry weight are reduced (by 80 and 53%, respectively), leaf and root growth are stimulated (by 41 and 86%, respectively). The difference in production between infected and uninfected millet plants can be accounted for by two processes: first, export of carbon to the parasite (accounting for 16% of the dry weight not gained); and second, parasite-induced reductions in host photosynthesis (accounting for 84% of the dry weight not gained). Striga hermonthica is dependent on carbon exported from the host, since the plant has low rates of photosynthesis coupled with high rates of respiration. The carbon balance model suggests that in mature S. hermonthica plants parasitic on millet, 85% of the carbon is host-derived. Carbon fluxes are also estimated using δ13C measurements, since S. hermonthica is a C3 plant parasitizing a C4 host. In conjunction with gas exchange measurements, these suggest that in root, stem and leaf of S. hermonthica, 87, 70 and 49% of carbon is hostderived, respectively.

Notes: In a study of the plant communities of two Australian rainforests, it was found that pioner species had high levels of nitrate reductase (EC 1.6.6.1) and were predominantly leaf nitrate assimilators. Under- and over-storey species had low levels of shoot and root nitrate reductase activity, and many of them showed little capacity for nitrate reduction even when nitrate ions were freely available. Although closed-forest species have lower levels of nitrate reductase than those of gaps and forest margins, their total nitrogen contents were similar, suggesting the former utilize nitrogen sources other than nitrate ions. Glutamine synthetase (EC 6.3.1.2) was present in the leaves of all species examined. In the leaves of pioneer species the chloroplastic isoform of glutamine synthetase predominted, while in most of the species typical of closed-forest the cytosolic isoform accounted for at least 40% of total leaf activity. Low levels of chloroplastic glutamine synthetase were correlated with a low capacity for leaf nitrate reduction, and both are characteristic of many species that regenerate and grow for some time in shade. Low levels of chloroplastic glutamine synthetase imply that, in some of these woody plants, photorespiratory ammonia is re-assimilated via cytosolic glutamine synthetase.

Notes: Photosynthesis and stomatal characteristics of the angiosperm parasite Striga her-monthica (Del.) Benth. have been compared with those of Antirrhinum majus L. cv. Suttons Yellow Rust Resistant, a related, non-parasitic species. The concentration of photosynthetic pigments in S. hermonthica leaves was less than 40% of those of A. majus leaves. Light saturated CO2 assimilation rate of S. hermonthica was less than 40% that of A. majus on a chlorophyll basis and under 20% than of A. majus on a leaf area basis.Stomata of Striga showed only partial closure in darkness, remained open in water stressed leaves and showed little response to exogenously applied abscisic acid. Stomatal conductance and transpiration were considerably higher in Striga compared with Antirrhinum. The high transpiration and low photosynthetic rates of S. hermonthica resulted in a low water use efficiency. The water relations of Striga leaves, while seemingly inappropriate for growth in drought prone environments, do appear to maximise nutrient and water acquisition from the host and as such may be an adaptation to the parasitic habit.

Notes: Samples of recently produced shoot material collected in winter/spring from common plant species of mulga vegetation in eastern and Western Australia were assayed for 13C and 15N natural abundance. 13C analyses showed only three of the 88 test species to exhibit C4 metabolism and only one of seven succulent species to be in CAM mode. Non-succulent winter ephemeral C3 species showed significantly lower mean δ13C values (– 28·0‰) than corresponding C3-type herbaceous perennials, woody shrubs or trees (– 26·9, – 25·7 and – 26·2‰, respectively), suggesting lower water stress and poorer water use efficiency in carbon acquisition by the former than latter groups of taxa. Corresponding values for δ15N of the above growth and life forms lay within the range 7·5–15·5‰. δ15N of soil NH4+ (mean 19·6‰) at a soft mulga site in Western Australia was considerably higher than that of NO3– (4·3‰). Shoot dry matter of Acacia spp. exhibited mean δ15N values (9·10 ± 0·6‰) identical to those of 37 companion non-N2-fixing woody shrubs and trees (9·06 ± 0·5‰). These data, with no evidence of nodulation, suggested little or no input of fixed N2 by the legumes in question. However, two acacias and two papilionoid legumes from a dune of wind-blown, heavily leached sand bordering a lake in mulga in Western Australia recorded δ15N values in the range 2·0–3·0‰ versus 6·4–10·7‰ for associated non-N2-fixing taxa. These differences in δ15N, and prolific nodulation of the legumes, indicated symbiotic inputs of fixed N in this unusual situation. δ15N signals of lichens, termites, ants and grasshoppers from mulga of Western Australia provided evidence of N2 fixation in certain termite colonies and by a cyanobacteria-containing species of lichen. Data are discussed in relation to earlier evidence of nitrophily and water availability constraints on nitrate utilization by mulga vegetation.

Notes: Analysis of soil solution from forest sites dominated by Eucalyptus grandis and Eucalyptus maculata indicates that soluble forms of organic nitrogen (amino acids and protein) are present in concentrations similar to those of mineral nitrogen (nitrate and ammonium). Experiments were conducted to determine the extent to which mycorrhizal associations might broaden nitrogen source utilization in Eucalyptus seedlings to include organic nitrogen. In isolation, species of ectomycorrhizal fungi from northern Australia show varying abilities to utilize mineral and organic forms of nitrogen as sole sources. Pisolithus sp. displayed strongest growth on NH4+, glutamine and asparagine, but grew poorly on protein, while Amanita sp. grew well both on mineral sources and on a range of organic sources (e.g. arginine, asparagine, glutamine and protein). In sterile culture, non-mycorrhizal seedlings of Eucalyptus grandis and Eucalyptus maculata grew well on mineral sources of nitrogen, but showed no ability to grow on sources of organic nitrogen other than glutamine. In contrast, mycorrhizal seedlings grew well on a range of organic nitrogen sources. These observations indicate that mycorrhizal associations confer on species of Eucalyptus the ability to broaden their resource base substantially with respect to nitrogen. This ability to utilize organic nitrogen was not directly related to that of the fungal symbiont in isolation. Seedlings mycorrhizal with Pisolithus sp. were able to assimilate sources of nitrogen (in particular histidine and protein) on which the fungus in pure culture appeared to grow weakly. Experiments in which plants were fed 15N-labelled ammonium were undertaken in order to investigate the influence of mycorrhizal colonization on the pathway of nitrogen metabolism. In roots and shoots of all seedlings, 15N was incorporated into the amide group of glutamine, and label was also found in the amino groups of glutamine, glutamic acid, γ-aminobutyric acid and alanine. Mycorrhizal colonization appeared to have no effect on the assimilation pathway and metabolism of [15N]H4+; labelling data were consistent with the operation of the glutamate synthase cycle in plants infected with either Pisolithus sp. (which in isolation assimilates via the glutamate synthase cycle) or Elaphomyces sp. (which assimilates via glutamate dehydrogenase). It is likely that the control of carbon supply to the mycorrhizal fungus from the host may have a profound effect on both the assimilatory pathway and the range of nitrogen sources that can be utilized by the association.

Notes: Chickpeas were grown with or without nitrate nitrogen feeding, or nodulated with Rhizobium leguminosarum. High [40°C day, 25°C night (HT)] and moderate [25°C day, 177°C night (LT)] temperature regimes were employed during growth. Growth rates, photosynthetic capacity and enzymes of carbon and nitrogen metabolism were monitored to assess the acclimatory capacity of the chickpea. Initial growth rates were stimulated by high temperatures, particularly in nitrate-fed and nodulated plants. Older HT plants had fewer laterals, smaller leaves, and fewer flowers were produced than in LT plants. There was some indication of an acclimation of photosynthesis to high temperatures and this was independent of nitrogen supply. Rubisco activity was increased by high growth temperatures. However, HT plants also had higher transpiration rates and lower water use efficiency than LT plants both in respective growth conditions and when compared in a common condition. High temperatures reduced shoot nitrate reductase activity but had little effect on root activity, which was the same if not greater than activity in LT roots. The amino acid, asparagine, was found at high concentrations in all treatments. Concentrations were maintained throughout growth in HT plants but declined with age in LT plants.

Notes: The amino acid compositions of the root xylem saps of Olax phyllanthi and a range of its common hosts were examined in native coastal heath in Western Australia and in pot cultures of Olax reliant on single hosts. When hosts specializing in the xylem transport of one major solute (asparagine, glutamine, histidine, arginine or proline) were exploited, the endophytic tissue of haustoria and the xylem sap of Olax showed much lower proportions of this than of other solutes, suggesting pronounced metabolic transformation prior to xylem loading by the parasite. However, the xylem sap of Olax did partly reflect the compositions of its hosts; for example, djenkolic acid and pipecolic acid were present when Olax was parasitic on species of Acacia, and levels of citrulline and aspartic acid were higher than normal when it exploited hosts transporting large amounts of these compounds. Back-flow of S-ethenyl cysteine, a novel amino acid specific to Olax, was observed to another root hemiparasite (Exocarpos sparteus) in native habitat and to certain non-parasitic hosts in water-stressed pot cultures. Haustoria exhibited high levels of glutamine synthetase but showed appreciable in vivo nitrate reductase activity only when on hosts with high xylem levels of nitrate.

Notes: A range of approaches was used to investigate how species within a fire-prone Banksia woodland in South West Australia exploited inorganic soil nitrogen sources and how this changes through the development of the fire chronosequence. Nitrate and ammonium were present in soil solution throughout the chronosequence but nitrate predominated in recently burnt sites. Mean shoot nitrate reductase activities were high for all species in recently burnt sites and showed little increase when nitrate was supplied via the transpiration stream. Nitrate reductase of shoots of most species was low at sites not burnt for several years, but following transpirational induction with nitrate, developed activities similar to those at recently burnt sites. The principal amino compounds transported in the xylem were species specific, including asparagine, glutamine and citrulline-dominated species, and changed little in relative composition across the chronosequence. Species most active in leaf nitrate reduction transported the largest amounts of nitrate in their xylem sap and proportional amounts of nitrate in xylem tended to be greatest in recently burnt sites. Most of the species examined appeared to be shoot rather than root nitrate assimilators, but marked differences were recorded in potential of leafy shoots of different species to reduce nitrate. As a general rule, shallow-rooted herbaceous, non-mycorrhizal or VAM-positive species had the highest capacity to reduce nitrate, whereas woody species with ericoid mycorrhizae or combined vesicular arbuscular/ectomycorrhizal associations exhibited little capacity to reduce nitrate in roots or shoots. It seems likely that this latter group utilize ammonium or even organic forms of nitrogen rather than nitrate. Some putative nitrogen-fixing species were active in reducing and transporting nitrate, others were virtually inactive in these respects.