Stress and Resilience in Combat-Related PTSD: Integration of Psychological Theory and Biological Mechanisms

Post-traumatic stress disorder (PTSD) is a debilitating psychiatric disorder that has a major impact on the ability to function effectively in daily life. PTSD may develop as a response to exposure to an event or events perceived as potentially harmful or life-threatening. It has high prevalence rates in the community, especially among vulnerable groups such as military personnel or those in emergency services. Despite extensive research in this field, the underlying mechanisms of the disorder remain largely unknown. The identification of risk factors for PTSD has posed a particular challenge as there can be delays in onset of the disorder, and most people who are exposed to traumatic events will not meet diagnostic criteria for PTSD. With the advent of the Diagnostic and Statistical Manual of Mental Disorders, Fifth Edition (DSM V), the classification for PTSD has changed from an anxiety disorder into the category of stress- and trauma-related disorders. This has the potential to refocus PTSD research on the nature of stress and the stress response relationship. This review focuses on some of the important findings from psychological and biological research based on early models of stress and resilience. Improving our understanding of PTSD by investigating both genetic and psychological risk and coping factors that influence stress response, as well as their interaction, may provide a basis for more effective and earlier intervention.

Agonosoma trilineatum (Heteroptera: Scutelleridae) a biological control agent of the weed bellyache bush, Jatropha gossypiifolia (Euphorbiaceae)

Bellyache bush, Jatropha gossypiifolia L., is a serious weed of northern Australia. Agonosoma trilineatum (F.) is an insect from tropical America released in Australia in 2003 as a biological control agent against bellyache bush. It feeds on seeds and has the potential to reduce seed production, thereby potentially reducing the rate of spread and recruitment. To test the host specificity of A. trilineatum, four biological responses to host plant species were determined: development of nymphs, oviposition preferences, adult feeding and frequency of mating. Development of nymphs to adults and adult feeding only occurred on three Jatropha spp. These species also supported mating and oogenesis but only J. gossypiifolia was accepted for oviposition. Mating did not occur in the presence of other plant species. The evidence indicates that there is little risk associated with the release of this insect species in Australia and probably other countries where this weed is a problem. The probability of this insect expanding its host range is low because multiple aspects of the biology would need to change simultaneously. A. trilineatum was released in Australia between 2003 and 2007. A Climex model indicated that coastal areas of Queensland and the Northern Territory would be climatically most suitable for this insect.

Evaluation of Ustilago sporoboli-indici as a classical biological control agent for invasive Sporobolus grasses in Australia

Sporobolus pyramidalis, S. africanus, S. natalensis, S. fertilis and S. jacquemontii, known collectively as the weedy Sporobolus grasses, are exotic weeds causing serious economic losses in grazing areas along Australia's entire eastern coast. In one of the first attempts to provide biological control for a grass, the potential of a smut, Ustilago sporoboli-indici, as a biological control agent for all five weedy Sporobolus spp. found in Australia was evaluated in glasshouse studies. Application of basidiospores to 21-day-old Sporobolus seedlings and subsequent incubation in a moist chamber (26 °C, 90% RH, 48 h) resulted in infection of S. pyramidalis, S. africanus, S. natalensis and S. fertilis but not S. jacquemontii. Host-range trials with 13 native Australian Sporobolus spp. resulted in infection of four native species. Evaluation of damage caused by the smut on two Australian native and two weedy Sporobolus spp. showed that the total numbers of flowers infected for the four grasses were in the following order: S. creber > S. fertilis > S. elongatus > S. natalensis with percentage flower infections of 21%, 14%, 12% and 3%, respectively. Significant differences (P = 0.001) were found when the numbers of infected flowers caused by each treatment were compared. The infection of the four native Sporobolus spp. by the smut indicated that it was not sufficiently host specific for release in Australia and the organism was rejected as a potential biological control agent. The implications of these results are discussed.

Prioritising potential guilds of specialist herbivores as biological control agents for prickly acacia through simulated herbivory

Understanding plant response to herbivory facilitates the prioritisation of guilds of specialist herbivores as biological control agents based on their potential impacts. Prickly acacia (Acacia nilotica ssp. indica) is a weed of national significance in Australia and is a target for biological control. Information on the susceptibility of prickly acacia to herbivory is limited, and there is no information available on the plant organ (i.e. leaf, shoot and root in isolation or in combination) most susceptible to herbivory. We evaluated the ability of prickly acacia seedlings, to respond to different types of simulated herbivory (defoliation, shoot damage, root damage and combinations), at varying frequencies (no herbivory, single, two and three events of herbivory) to identify the type and frequency of herbivory that will be required to reduce the growth and vigour. Defoliation and shoot damage, individually, had a significant negative impact on prickly acacia seedlings. For the defoliation to be effective, more than two defoliation events were required, whereas a single bout of shoot damage was enough to cause a significant reduction in plant vigour. A combination of defoliation + shoot damage had the greatest negative impact. The study highlights the need to prioritise specialist leaf and shoot herbivores as potential biological control agents for prickly acacia.

New efforts at biological control of Mikania micrantha H.B.K. (Asteraceae) in Papua New Guinea and Fiji.

New efforts at biological control of Miconia calvescens (Melastomataceae) is a serious invader in the tropical Pacific, including the Hawaiian and Tahitian Islands, and currently poses a major threat to native biodiversity in the Wet Tropics of Australia. The species is fleshy-fruited, small-seeded and shade tolerant, and thus has the potential to be dispersed widely and recruit in relatively intact rainforest habitats, displacing native species. Understanding and predicting the rate of spread is critical for the design and implementation of effective management actions. We used an individual-based model incorporating a dispersal function derived from dispersal curves for similar berry-fruited native species, and life-history parameters of fecundity and mortality to predict the spatial structure of a Miconia population after a 30 year time period. We compared the modelled population spatial structure to that of an actual infestation in the rainforests of north Queensland. Our goal was to assess how well the model predicts actual dispersion and to identify potential barriers and conduits to seed movement and seedling establishment. The model overpredicts overall population size and the spatial extent of the actual infestation, predicting individuals to occur at a maximum 1,750 m from the source compared with the maximum distance of any detected individual in the actual infestation of 1,191 m. We identify several characteristic features of managed invasive populations that make comparisons between modelled outcomes and actual infestations difficult. Our results suggest that the model’s ability to predict both spatial structure and spread of the population will be improved by incorporating a spatially explicit element, with dispersal and recruitment probabilities that reflect the relative suitability of different parts of the landscape for these processes. Mikania micrantha H.B.K. (Asteraceae) in Papua New Guinea and Fiji.

Prospects for the biological control of the weedy sporobolus grasses in Australia.

Bill Palmer and colleagues recently published their paper 'Prospects for the biological control of the weedy sporobolus grasses in Australia' in Proceedings of the 16th Australian Weeds Conference. The paper gives a summary of a recent project to find a biological control for the weedy sporobolus grasses, which include giant rat's tail grass. Southern Africa was surveyed for potential agents and two, a leaf smut and a stem wasp, were selected for follow up studies. Unfortunately, they could not rear the stem wasp in the laboratory and the leaf smut infected four of the Australian native Sporobolus spp. and was therefore rejected. This project was one of the first attempts at biological control of a grass.

Applications of CLIMEX modelling leading to improved biological control

Wilmot Senaratne, Bill Palmer and Bob Sutherst recently published their paper 'Applications of CLIMEX modelling leading to improved biological control' in Proceedings of the 16th Australian Weeds Conference. They looked at three examples where modern climate matching techniques using computer software produces decisions and results than might happen using previous techniques such as climadiagrams. Assessment of climatic suitability is important at various stages of a biological control project; from initial foreign exploration, to risk assessment in preparation for the release of a particular agent, through to selection of release sites that maximise the agent´s chances of initial establishment. It is now also necessary to predict potential future distributions of both target weeds and agents under climate change.

Factors influencing the release and establishment of weed biological control agents

Mike Day and colleagues recently published their paper 'Factors influencing the release and establishment of weed biocontrol agents' in Proceedings of the 16th Australian Weeds Conference. The CRC for Australian Weed Management facilitated an investigation into the factors influencing the release and establishment of weed biological control agents on a wide variety of Australian weeds. The investigation improved the understanding of post-release ecology of biocontrol agents and generated recommendations for best practice. Factors affecting successful establishment on the weed include host plant characteristics, size of releases, dispersal power of the agent, predation and parasitism, and climate. A best practice guide was produced by the CRC to assist practitioners in designing robust release strategies to increase rates of establishment.

Pre-release evaluation of the efficacy of the leaf-sucking bug Carvalhotingis visenda (Heteroptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae)

In classical weed biological control, assessing weed response to simulated herbivory is one option to assist in the prioritization of available agents and prediction of their potential efficacy. Previously reported simulated herbivory studies suggested that a specialist herbivore in the leaf-feeding guild is desirable as an effective biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae), an environmental weed that is currently a target for biological control. In this study, we tested (i) whether the results from glasshouse-based simulated herbivory can be used to prioritise potential biological control agents by evaluating the impact of a leaf-sucking tingid bug Carvalhotingis visenda (Drake & Hambleton) (Hemiptera: Tingidae) in quarantine; and (ii) the likely effectiveness of low- and high-densities of the leaf-sucking tingid after its release in the field. The results suggest that a single generation of C. visenda has the potential to reduce leaf chlorophyll content significantly, resulting in reduced plant height and leaf biomass. However, the impact of one generation of tingid herbivory on below-ground plant components, including the roots and tuber size and biomass, were not significant. These findings are consistent with results obtained from a simulated herbivory trial, highlighting the potential role of simulated herbivory studies in agent prioritisation.

The leaf-tying moth Hypocosmia pyrochroma (Lep., Pyralidae), a host-specific biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

Cat's claw creeper, Macfadyena unguis-cati, a major environmental weed in coastal and sub-coastal areas of Queensland and New South Wales, Australia is a target for classical biological control. Host specificity of Hypocosmia pyrochroma Jones (Lep., Pyralidae), as a potential biological control agent was evaluated on the basis of no-choice and choice larval feeding and survival, and adult oviposition preference tests, involving 38 plant species in 10 families. In no-choice tests, larval feeding and development occurred only on cat's claw creeper. In choice tests, oviposition and larval development was evident only on cat's claw creeper. The results support the host-specificity tests conducted in South Africa, and suggest that H. pyrochroma is a highly specific biological control agent that does not pose any risk to non-target plants tested in Australia. This agent has been approved for field release by relevant regulatory authorities in Australia.

Will biological control of Lantana camara ever succeed? Patterns, processes & prospects

Despite biocontrol research spanning over 100 years, the hybrid weed, commonly referred to as Lantana camara, is not under adequate control. Host specificity and varietal preference of released agents, climatic suitability of a region for released agents, number of agents introduced and range or area of infestation appear to play a role in limiting biocontrol success. At least one of 41 species of mainly leaf- or flower-feeding insects has been introduced, or spread, to 41 of the 70 countries or regions where lantana occurs. Over half (26) of these species have established, achieving varying levels of herbivory and presumably some degree of control. Accurate taxonomy of the plant and adaptation of potential agents to the host plant are some of the better predictors of at least establishment success. Retrospective analysis of the hosts of introduced biocontrol agents for L. camara show that a greater proportion of agents that were collected from L. camara or Lantana urticifolia established, than agents that were collected from other species of Lantana. Of the introduced agents that had established and were oligophagous, 18 out of 22 established. The proportion of species establishing, declined with the number of species introduced. However, there was no trend when oceanic islands were treated separately from mainland areas and the result is likely an artefact of how introductions have changed over time. A calculated index of the degree of herbivory due to agents known to have caused some damage per country, was not related to land area infested with lantana for mainlands nor for oceanic islands. However, the degree of herbivory is much higher on islands than mainlands. This difference between island and mainland situations may reflect population dynamics in patchy or metapopulation landscapes. Basic systematic studies of the host remain crucial to successful biocontrol, especially of hybrid weeds like L. camara. Potential biocontrol agents should be monophages collected from the most closely related species to the target weed or be phytophages that attack several species of lantana. Suitable agents should be released in the most ideal ecoclimatic area. Since collection of biocontrol agents has been limited to a fraction of the known number of phytophagous species available, biocontrol may be improved by targeting insects that feed on stems and roots, as well as the agents that feed on leaves and flowers.

Predicting risk and benefit a priori in biological control of invasive plant species: a systems modelling approach

While the method using specialist herbivores in managing invasive plants (classical biological control) is regarded as relatively safe and cost-effective in comparison to other methods of management, the rarity of strict monophagy among insect herbivores illustrates that, like any management option, biological control is not risk-free. The challenge for classical biological control is therefore to predict risks and benefits a priori. In this study we develop a simulation model that may aid in this process. We use this model to predict the risks and benefits of introducing the chrysomelid beetle Charidotis auroguttata to manage the invasive liana Macfadyena unguis-cati in Australia. Preliminary host-specificity testing of this herbivore indicated that there was limited feeding on a non-target plant, although the non-target was only able to sustain some transitions of the life cycle of the herbivore. The model includes herbivore, target and non-target life history and incorporates spillover dynamics of populations of this herbivore from the target to the non-target under a variety of scenarios. Data from studies of this herbivore in the native range and under quarantine were used to parameterize the model and predict the relative risks and benefits of this herbivore when the target and non-target plants co-occur. Key model outputs include population dynamics on target (apparent benefit) and non-target (apparent risk) and fitness consequences to the target (actual benefit) and non-target plant (actual risk) of herbivore damage. The model predicted that risk to the non-target became unacceptable (i.e. significant negative effects on fitness) when the ratio of target to non-target in a given patch ranged from 1:1 to 3:2. By comparing the current known distribution of the non-target and the predicted distribution of the target we were able to identify regions in Australia where the agent may be pose an unacceptable risk. By considering risk and benefit simultaneously, we highlight how such a simulation modelling approach can assist scientists and regulators in making more objective decisions a priori, on the value of releasing specialist herbivores as biological control agents.

Specificity of Carvalhotingis visenda (Hemiptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

Biological control is considered the most suitable management option for cat's claw creeper, Macfadyena unguis-cati, a major environmental weed in coastal and sub-coastal areas of Queensland and New South Wales, Australia. The potential host range of the leaf-sucking bug, Carvalhotingis visenda (Hemiptera: Tingidae) was evaluated on the basis of nymphal survival and development, adult feeding and survival, and oviposition preference using choice and no-choice tests involving 38 plant species in 10 families. In no-choice tests, although adults survived on a few of the non-target plants, no eggs were laid on any of the non-target plants. In no-choice condition, the tingid oviposits and completes nymphal development only on M. unguis-cati. There was also no visible feeding damage on any of the non-target plants. In choice tests, adults showed distinct preference for M. unguis-cati, and the preference level increased over time as the tingids moved away from the non-target plants. At the end of the trial no adults were evident on any of the non-target plants. Host specificity tests confirm that the tingid is a highly host specific biocontrol agent, and does not pose risk to any non-target plants in Australia. This agent has been approved for field release by the relevant regulatory authorities in Australia.