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The grazing lands of northern Australia contain a substantial soil organic carbon (SOC) stock due to the large land area. Manipulating SOC stocks through grazing management has been presented as an option to offset national greenhouse gas emissions from agriculture and other industries. However, research into the response of SOC stocks to a range of management activities has variously shown positive, negative or negligible change. This uncertainty in predicting change in SOC stocks represents high project risk for government and industry in relation to SOC sequestration programs. In this paper, we seek to address the uncertainty in SOC stock prediction by assessing relationships between SOC stocks and grazing land condition indicators. We reviewed the literature to identify land condition indicators for analysis and tested relationships between identified land condition indicators and SOC stock using data from a paired-site sampling experiment (10 sites). We subsequently collated SOC stock datasets at two scales (quadrat and paddock) from across northern Australia (329 sites) to compare with the findings of the paired-site sampling experiment with the aim of identifying the land condition indicators that had the strongest relationship with SOC stock. The land condition indicators most closely correlated with SOC stocks across datasets and analysis scales were tree basal area, tree canopy cover, ground cover, pasture biomass and the density of perennial grass tussocks. In combination with soil type, these indicators accounted for up to 42% of the variation in the residuals after climate effects were removed. However, we found that responses often interacted with soil type, adding complexity and increasing the uncertainty associated with predicting SOC stock change at any particular location. We recommend that caution be exercised when considering SOC offset projects in northern Australian grazing lands due to the risk of incorrectly predicting changes in SOC stocks with change in land condition indicators and management activities for a particular paddock or property. Despite the uncertainty for generating SOC sequestration income, undertaking management activities to improve land condition is likely to have desirable complementary benefits such as improving productivity and profitability as well as reducing adverse environmental impact.

Soil and land-management interactions in Australian native-forest regrowth remain a major source of uncertainty in the context of the global carbon economy. We sampled soil total organic C (TOC) and soil total N (TN) stocks at 45 sites within the Brigalow ecological community of the Brigalow Belt bioregion, Queensland, Australia. The sites were matched as triplets representing three land uses, specifically: uncleared native brigalow forest (‘Remnant’); grassland pasture (‘Pasture’), derived by clearing native vegetation and maintained as pasture for a minimum of 10 years, and; regrowing native brigalow forest (‘Regrowth’, stand ages ranging from 10 to 58 years) that had developed spontaneously after past vegetation clearing for pasture establishment. Soil TOC fractions and natural abundance of soil C and N isotopes were examined to obtain insight into C and N dynamics. An updated above- and belowground carbon budget for the bioregions was generated. Average soil TOC stocks at 0–0.3-m depth ranged from 19 to 79Mgha-1 and soil TN stocks from 1.8 to 7.1Mgha-1 (2.5th and 97.5th percentiles, respectively). A trend in stocks was apparent with land use: Remnant>Regrowth ~ Pasture sites. Soil δ13C ranged from –14 to –27‰, and soil δ15N ranged from 4‰ to 17‰, in general reflecting the difference between Pasture (C4-dominated) land use and N2-fixing (C3-dominated) Remnant and Regrowth. Mid-infrared spectroscopy predicted C fractions as a percentage of soil TOC stock, which ranged from 5% to 60% (particulate), 20–80% (humus) and 9–30% (resistant/inert). The geo-referenced soil and management information we collected is important for the calibration of C models, for the estimation of national C accounts, and to inform policy developments in relation to land-resource management undertaken within the Brigalow Belt bioregions of Australia.

Beef cattle grazing is the dominant land use in the extensive tropical and sub-tropical rangelands of northern Australia. Despite the considerable knowledge on land and herd management gained from both research and practical experience, the adoption of improved management is limited by an inability to predict how changes in practices and combinations of practices will affect cattle production, economic returns and resource condition. To address these issues, past Australian and international research relating to four management factors that affect productivity and resource condition was reviewed in order to identify key management principles. The four management factors considered were stocking rates, pasture resting, prescribed fire, and fencing and water point development for managing grazing distribution. Four management principles for sound grazing management in northern Australia were formulated as follows: (1) manage stocking rates to meet goals for livestock production and land condition; (2) rest pastures to maintain them in good condition or to restore them from poor condition to increase pasture productivity; (3) devise and apply fire regimes that enhance the condition of grazing land and livestock productivity while minimising undesirable impacts; and (4) use fencing and water points to manipulate grazing distribution. Each principle is supported by several more specific guidelines. These principles and guidelines, and the supporting research on which they are based, are presented.

This study aimed to unravel the effects of climate, topography, soil, and grazing management on soil organic carbon (SOC) stocks in the grazing lands of north-eastern Australia. We sampled for SOC stocks at 98 sites from 18 grazing properties across Queensland, Australia. These samples covered four nominal grazing management classes (Continuous, Rotational, Cell, and Exclosure), eight broad soil types, and a strong tropical to subtropical climatic gradient. Temperature and vapour-pressure deficit explained >80% of the variability of SOC stocks at cumulative equivalent mineral masses nominally representing 0–0.1 and 0–0.3m depths. Once detrended of climatic effects, SOC stocks were strongly influenced by total standing dry matter, soil type, and the dominant grass species. At 0–0.3m depth only, there was a weak negative association between stocking rate and climate-detrended SOC stocks, and Cell grazing was associated with smaller SOC stocks than Continuous grazing and Exclosure. In future, collection of quantitative information on stocking intensity, frequency, and duration may help to improve understanding of the effect of grazing management on SOC stocks. Further exploration of the links between grazing management and above- and below-ground biomass, perhaps inferred through remote sensing and/or simulation modelling, may assist large-area mapping of SOC stocks in northern Australia.

The emerging carbon economy will have a major impact on grazing businesses because of significant livestock methane and land-use change emissions. Livestock methane emissions alone account for ~11% of Australia's reported greenhouse gas emissions. Grazing businesses need to develop an understanding of their greenhouse gas impact and be able to assess the impact of alternative management options. This paper attempts to generate a greenhouse gas budget for two scenarios using a spread sheet model.The first scenario was based on one land-type ‘20-year-old brigalow regrowth' in the brigalow bioregion of southern-central Queensland. The 50 year analysis demonstrated the substantially different greenhouse gas outcomes and livestock carrying capacity for three alternative regrowth management options: retain regrowth (sequester 71.5 t carbon dioxide equivalents per hectare, CO2-e/ha), clear all regrowth (emit 42.8 t CO2-e/ha) and clear regrowth strips (emit 5.8 t CO2-e/ha). The second scenario was based on a ‘remnant eucalypt savanna-woodland' land type in the Einasleigh Uplands bioregion of north Queensland. The four alternative vegetation management options were: retain current woodland structure (emit 7.4 t CO2-e/ha), allow woodland to thicken increasing tree basal area (sequester 20.7 t CO2-e/ha), thin trees less than 10cm diameter (emit 8.9 t CO2-e/ha), and thin trees <20cm diameter (emit 12.4 t CO2-e/ha).Significant assumptions were required to complete the budgets due to gaps in current knowledge on the response of woody vegetation, soil carbon and non-CO2soil emissions to management options and land-type at the property scale. The analyses indicate that there is scope for grazing businesses to choose alternative management options to influence their greenhouse gas budget. However, a key assumption is that accumulation of carbon or avoidance of emissions somewhere on a grazing business (e.g. in woody vegetation or soil) will be recognised as an offset for emissions elsewhere in the business (e.g. livestock methane). This issue will be a challenge for livestock industries and policy makers to work through in the coming years.

The Notch signalling pathway regulates cell fate choices during both vertebrate and invertebrate development. In the Drosophila wing disc, the activation of Notch by its ligands Delta and Serrate is required to make the dorsoventral boundary, where several genes, such as wingless and cut, are expressed in a 2- to 4-cell-wide domain. The interactions between Notch and its ligands are modulated by Fringe via a mechanism that may involve post-transcriptional modifications of Notch. The ligands themselves also help to restrict Notch activity to the dorsoventral boundary cells, because they antagonise the activation of the receptor in the cells where their expression is high. This function of the ligands is critical to establish the polarity of signalling, but very little is known about the mechanisms involved in the interactions between Notch and its ligands that result in suppression of Notch activity. The extracellular domain of Notch contains an array of 36 EGF repeats, two of which, repeats 11 and 12, are necessary for direct interactions between Notch with Delta and Serrate. We investigate here the function of a region of the Notch extracellular domain where several missense mutations, called Abruptex, are localised. These Notch alleles are characterised by phenotypes opposite to the loss of Notch function and also by complex complementation patterns. We find that, in Abruptex mutant discs, only the negative effects of the ligands and Fringe are affected, resulting in the failure to restrict the expression of cut and wingless to the dorsoventral boundary. We suggest that Abruptex alleles identify a domain in the Notch protein that mediates the interactions between Notch, its ligands and Fringe that result in suppression of Notch activity.

The legs of Drosophila are divided into segments along the proximodistal axis by flexible structures called joints. The separation between segments is already visible in the imaginal disc as folds of the epithelium, and cells at segment boundaries have different morphology during pupal development. We find that Notch is locally activated in distal cells of each segment, as demonstrated by the restricted expression of the Enhancer of split mbeta gene, and is required for the formation of normal joints. The genes fringe, Delta, Serrate and Suppressor of Hairless, also participate in Notch function during leg development, and their expression is localised within the leg segments with respect to segment boundaries. The failure to form joints when Notch signalling is compromised leads to shortened legs, suggesting that the correct specification of segment boundaries is critical for normal leg growth. The requirement for Notch during leg development resembles that seen during somite formation in vertebrates and at the dorsal ventral boundary of the wing, suggesting that the creation of boundaries of gene expression through Notch activation plays a conserved role in co-ordinating growth and patterning.

A common consequence of Notch signalling in Drosophila is the transcriptional activation of seven Enhancer of split [E(spl)] genes, which encode a family of closely related basic-helix-loop-helix transcriptional repressors. Different E(spl) proteins can functionally substitute for each other, hampering loss-of-function genetic analysis and raising the question of whether any specialization exists within the family. We expressed each individual E(spl) gene using the GAL4-UAS system in order to analyse their effect in a number of cell fate decisions taking place in the wing imaginal disk. We focussed on sensory organ precursor determination, wing vein determination and wing pattern formation. All of the E(spl) proteins affect the first two processes in the same way, namely they antagonize neural precursor and vein fates. Yet, the efficacy of this antagonism is quite distinct: E(spl)mbeta has the strongest vein suppression effect, whereas E(spl)m8 and E(spl)m7 are the most active bristle suppressors. During wing patterning, Notch activity orchestrates a complex sequence of events that define the dorsoventral boundary of the wing. We have discerned two phases within this process based on the sensitivity of N loss-of-function phenotypes to concomitant expression of E(spl) genes. E(spl) proteins are initially involved in repression of the vg quadrant enhancer, whereas later they appear to relay the Notch signal that triggers activation of cut expression. Of the seven proteins, E(spl)mgamma is most active in both of these processes. In conclusion, E(spl) proteins have partially redundant functions, yet they have evolved distinct preferences in implementing different cell fate decisions, which closely match their individual normal expression patterns.

A cis‐acting sequence from the Drosophila melanogaster dopa decarboxylase (Ddc) gene is selectively required for Ddc expression in the central nervous system. We analyze several parameters influencing the function of the sequence element and describe a factor which interacts with it and mediates CNS expression of Ddc. The element, element I, can function in vivo when included on a synthetic oligonucleotide inserted near its normal location, or closer to the RNA startpoint. It displays partial activity when inverted. Two different 2‐bp mutations in element I abolish its ability to stimulate neuronal Ddc expression in the CNS. A factor present in embryonic nuclear extracts specifically protects element I in DNase I footprinting assays. The binding affinity of this factor is reduced by each alteration of element I that inhibits neuronal expression, indicating a role in mediating CNS expression of Ddc. Element I alone has no detectable activity when placed adjacent to a heterologous promoter, although 2.2 kb of 5′ Ddc sequences direct correct cell‐specific expression of a heterologous promoter.

Abstract: The potential of Pontoscolex corethrurus to repair the physical degradation of a compacted Oxisol was studied. The Oxisol from Kingaroy, Queensland, Australia was uniformly packed to four treatments of different bulk densities (0.95, 1.15, 1.25, 1.35 Mg m–3) in pots of 0.24 m diameter and 0.22 m deep. Each pot was inoculated with 12 earthworms (equivalent to 300 m–2) and maintained close to field capacity water content for 3 months, after which selected soil physical (dry bulk density, penetration resistance, water infiltration), worm activity (cast production, worm weight) measurements and image analysis were carried out. Results showed that worm numbers were maintained at the initial levels in all the treatments except in the 1.35 Mg m–3 treatment, where there was a 33% decrease. The weight of surface casts per surviving worm was the greatest in the 1.35 Mg m–3 treatment compared to the lower density treatments. Final soil density was lessened in all treatments to a depth of 0.2 m. Surface cast production was positively correlated with the percentage reduction in bulk density. The greatest percentage reduction in bulk density was in the 1.35 Mg m–3 treatment and was equivalent to a doubling of soil aeration (to 18.4%). Penetration resistance to the 0.2-m layer was also reduced and values were less than 2 MPa. Slumping of the surface soil was evident in both the lowest and the highest bulk densities resulting in low rates of water infiltration. Image analysis supported the soil physical properties showing an abundance of both fine pores and worm channels in the three lowest bulk densities, and lesser activity (concentrated in the immediate soil surface) in the 1.35 Mg m–3 treatment.

The effects on beef cattle growth performance and carcass characteristics of feeding silages produced by altered fermentations were determined. Alpha-amylase was added at 0 or .05% (wet basis) and sorbic acid was added at 0 or .10% (wet basis) to chopped whole corn plants before ensiling in a 2 × 2 factorial arrangement of treatments. For three successive years, 40 beef heifers (224 kg) were fed these silages for 80 d, finished on corn-and-cob meal (107 d) and slaughtered when backfat thickness over the 13th rib reached 12 mm. Silages treated with α-amylase had a slightly higher percentage of N-free extract (P< .10). Silages treated with sorbic acid had lower percentages of ADF N (P<.10). During the silage-feeding phase, heifers fed silages treated with α-amylase gained more (P< .01) daily than heifers fed the other two silages (.84 vs .78 kg) and they were more efficient (P < .01) in weight gain per unit of dry feed consumed (.149 vs .139 kg). During the finishing phase, heifers that previously had been fed the α-amylase-treated silage continued to have higher (P< .05) ADG (.93 vs .87 kg), although all were fed the same diet during this period. Added sorbic acid had no effect on ADG in either period. The percentage of kidney fat in heifers on the α-amylase treatments was increased (P< .02, 2.2 vs 2.0). The biological mechanisms associated with the beneficial results of α-amylase addition are not understood yet.

Grainyhead belongs to a recently identified group of transcription factors which share a 250-amino-acid domain required for binding to DNA and a carboxy-terminal dimerization domain. The activities of Grainyhead and other members of the family appear to be modulated so that they can participate in different developmental processes. We have examined the structure and function of mRNAs from the Drosophila grainyhead gene and demonstrated that alternate splicing is responsible for generating a neuroblast-specific isoform of the protein. A mutation which abolishes this isoform results in pupal and adult lethality. Reporter genes containing different Grainyhead binding sites exhibit tissue-specific patterns of expression that correlate with the Grainyhead isoforms, suggesting that the alternate splicing serves to alter the repertoire of target genes controlled in the neuroblasts.

The Enhancer of split locus is required during many cell-fate decisions in Drosophila, including the segregation of neural precursors in the embryo. We have generated monoclonal antibodies that recognise some of the basic helix-loop-helix proteins encoded by the Enhancer of split locus and have used them to examine expression of Enhancer of split proteins during neurogenesis. The proteins are expressed in a dynamic pattern in the ventral neurogenic region and are confined to those ectodermal cells that surround a neuroblast in the process of delaminating. There is no staining in the neuroblasts themselves. We have also examined the relationship between Enhancer of split protein accumulation and the Notch signalling pathway. Protein expression is abolished in a number of neurogenic mutant backgrounds, including Notch, but is increased as a result of expressing a constitutively active Notch product. We conclude that Notch signalling activity is directly responsible for the accumulation of basic helix-loop-helix proteins encoded by the Enhancer of split locus.

The basic helix-loop-helix proteins of the Enhancer of split complex constitute a link between activation of the transmembrane receptor Notch and the resulting effects on transcription of downstream genes. The Suppressor of Hairless protein is the intermediary between Notch activation and expression of all Enhancer of split genes even though individual genes have distinct patterns of expression in imaginal discs. A comparison between the phenotypes produced by Notch, Suppressor of Hairless and Enhancer of split mutations in the wing and thorax indicate that Suppressor of Hairless and Notch requirements are indistinguishable, but that Enhancer of split activity is only essential for a subset of developmental processes involving Notch function. Likewise, the ectopic expression of Enhancer of split proteins does not reproduce all the consequences typical of ectopic Notch activation. We suggest that the Notch pathway bifurcates after the activation of Suppressor of Hairless and that Enhancer of split activity is not required when the consequence of Notch function is the transcriptional activation of downstream genes. Transcriptional activation mediated by Suppressor of Hairless and transcriptional repression mediated by Enhancer of split could provide greater diversity in the response of individual genes to Notch activity.

The cells along the dorsoventral boundary of the Drosophila wing imaginal disc have distinctive properties and their specification requires Notch activity. Later in development, these cells will form the wing margin, where sensory organs and specialised trichomes appear in a characteristic pattern. We find that Notch is locally activated in these cells, as demonstrated by the restricted expression of the Enhancer of split proteins in dorsal and ventral cells abutting the D/V boundary throughout the third larval instar. Furthermore other genes identified by their involvement in Notch signaling during neurogenesis, such as Delta and Suppressor of Hairless, also participate in Notch function at the dorsoventral boundary. In addition, Serrate, a similar transmembrane protein to Delta, behaves as a ligand required in dorsal cells to activate Notch at the boundary. Notch gain-of-function alleles in which Notch activity is not restricted to the dorsoventral boundary cause miss-expression of cut and wingless and overgrowth of the disc, illustrating the importance of localised Notch activation for wing development.

Notch function is required at the dorsoventral boundary of the developing Drosophila wing for its normal growth and patterning. We find that clones of cells expressing either Notch or its ligands Delta and Serrate in the wing mimic Notch activation at the dorsoventral boundary producing non-autonomous effects on proliferation, and activating expression of the target genes E(spl), wingless and cut. The analysis of these clones reveals several mechanisms important for maintaining and delimiting Notch function at the dorsoventral boundary. First, Notch activation in the wing leads to increased production of Delta and Serrate generating a positive feedback loop that maintains signalling. We propose that during normal development, wingless co-operates with Notch to reinforce this positive feedback and Cut, which is activated by Notch at late stages, acts antagonistically to prevent Delta and Serrate expression. Second, high levels of Delta and Serrate have a dominant negative effect on Notch, so that at late stages Notch can only be activated in cells next to the ligand-producing cells. Thus the combined effects of Notch and its target genes cut and wingless regulate the expression of Notch ligands which restrict Notch activity to the dorsoventral boundary.

The veins in the Drosophila wing have a characteristic width, which is regulated by the activity of the Notch pathway. The expression of the Notch-ligand Delta is restricted to the developing veins, and coincides with places where Notch transcription is lower. We find that this asymmetrical distribution of ligand and receptor leads to activation of Notch on both sides of each vein within a territory of Delta-expressing cells, and to the establishment of boundary cells that separate the vein from adjacent interveins. In these cells, the expression of the Enhancer of split gene m beta is activated and the transcription of the vein-promoting gene veinlet is repressed, thus restricting vein differentiation. We propose that the establishment of vein thickness utilises a combination of mechanisms that include: (1) independent regulation of Notch and Delta expression in intervein and vein territories, (2) Notch activation by Delta in cells where Notch and Delta expression overlaps, (3) positive feedback on Notch transcription in cells where Notch has been activated and (4) repression of veinlet transcription by E(spl)m beta and maintenance of Delta expression by veinlet/torpedo activity.

The Notch signaling pathway is known to regulate cell fate decisions in a variety of organisms from worms to humans. Although several components of the pathway have been characterized, the actual mechanism and molecular results of signaling remain elusive. We have examined the role of the Notch signaling pathway in the transcriptional regulation of two Drosophila Enhancer of split [E(spl)] genes, whose gene products have been shown to be downstream players in the pathway. Using a reporter assay system in Drosophila tissue culture cells, we have observed a significant induction of E(spl) m gamma and m delta expression after cotransfection with activated Notch. Characterization of the 5' regulatory regions of these two genes led to the identification of a number of target sites for the Suppressor of Hairless [Su(H)] protein, a transcription factor activated by Notch signaling. We show that Notch-inducible expression of E(spl) m gamma and m delta both in cultured cells and in vivo is dependent on functional Su(H). Although overexpression of Su(H) augments the level of induction of the reporter genes by activated Notch, Su(H) alone is insufficient to produce high levels of transcriptional activation. Despite the synergy observed between activated Notch and Su(H), the former affects neither the nuclear localization nor the DNA binding activity of the latter.

We have mapped the regions in the Drosophila melanogaster tissue-specific transcription factor Grainyhead that are required for DNA binding and dimerization. These functional domains correspond to regions conserved between Grainyhead and the vertebrate transcription factor CP2, which we show has similar activities. The identified DNA-binding domain is large (263 amino acids) but contains a smaller core that is able to interact with DNA at approximately 400-fold lower affinity. The major dimerization domain is located in a separate region of the protein and is required to stabilize the interactions with DNA. Our data also suggest that Grainyhead activity can be modulated by an N-terminal inhibitory domain.

In both clam oocytes and sea urchin eggs, fertilization triggers the synthesis of a set of proteins specified by stored maternal mRNAs. One of the most abundant of these (p41) has a molecular weight of 41,000. This paper describes the identification of p41 as the small subunit of ribonucleotide reductase, the enzyme that provides the precursors necessary for DNA synthesis. This identification is based mainly on the amino acid sequence deduced from cDNA clones corresponding to p41, which shows homology with a gene in Herpes Simplex virus that is thought to encode the small subunit of viral ribonucleotide reductase. Comparison with the B2 (small) subunit of Escherichia coli ribonucleotide reductase also shows striking homology in certain conserved regions of the molecule. However, our attention was originally drawn to protein p41 because it was specifically retained by an affinity column bearing the monoclonal antibody YL 1/2, which reacts with alpha-tubulin (Kilmartin, J. V., B. Wright, and C. Milstein, 1982, J. Cell Biol., 93:576-582). The finding that this antibody inhibits the activity of sea urchin embryo ribonucleotide reductase confirmed the identity of p41 as the small subunit. The unexpected binding of the small subunit of ribonucleotide reductase can be accounted for by its carboxy-terminal sequence, which matches the specificity requirements of YL 1/2 as determined by Wehland et al. (Wehland, J., H. C. Schroeder, and K. Weber, 1984, EMBO [Eur. Mol. Biol. Organ.] J., 3:1295-1300). Unlike the small subunit, there is no sign of synthesis of a corresponding large subunit of ribonucleotide reductase after fertilization. Since most enzymes of this type require two subunits for activity, we suspect that the unfertilized oocytes contain a stockpile of large subunits ready for combination with newly made small subunits. Thus, synthesis of the small subunit of ribonucleotide reductase represents a very clear example of the developmental regulation of enzyme activity by control of gene expression at the level of translation.