Work Package 1.3 - Biodiversity and Ecosystems
Researchers at the James Hutton Institute, the Royal Botanic Garden Edinburgh and Biomathematics and Statistics Scotland are working together to provide scientific evidence to underpin the Scottish Biodiversity Strategy. At the core of our work is understanding how management actions interact with biodiversity to deliver the outcomes we want. This ranges from understanding how the genetic diversity in one crop can be exploited to make farming more resilient to restoring populations of rare mountain plants. In addition, we use this understanding to provide indicators of how healthy ecosystems are for use by policy makers.
Aim of Research
There remain large gaps in understanding how changes in management and the environment alter community composition and, hence, ecosystem functions. This Research Deliverable (RD) addresses how ecosystem functions are regulated by the traits of species present, and how potential limits for the maintenance of ecosystem function can be captured in ecosystem health metrics. It also has the aim of identifying indices of ecosystem function that can be applied to assess ecosystem health at a range of scales (from field to national) so that management actions can be targeted to improve ecosystem health and to conserve and restore biodiversity.
Our work on crop mixtures has resulted in a paper in a special feature in Journal of Ecology on facilitative mechanisms underlying the effects of biodiversity on ecosystem functioning where we synthesise the links between ecological theory and agronomic performance of species and cultivar mixtures. In addition to improving outputs by increasing crop diversity at the field scale, a range of publications have identified the food security, health and environmental benefits of increasing crop diversity at the farm and landscape scale, particularly through the increased use of legume crops in Scottish farming. We have continued to build genetic stock and expertise in propagation and replanting of Cicerbita alpina (alpine sow thistle) and have used this material to build up populations in the Cairngorms. Similarly, we are starting to build up stock of Woodsia ilvensis (oblong woodsia). We have continued to explore the potential for Bere barley to supply genetic material for breeding to improve phosphorus and manganese acquisition and heavy metal tolerance and published a number of papers on this. The Glen Finglas research has produced two papers, one shows that the slow pace of change in vegetation composition is having little functional impact where grazing has been increased or abandoned, whilst the other has shown that bird species composition is starting to change towards more woodland and scrub species where succession to woodland is starting. We have provided a number of SEFARI case studies including one on the issues of grazing in the uplands based on the Glen Finglas work and another on the impact of liming on biodiversity which shows that hill grasslands are relatively resilient to increased liming and there is little impact on plants worms or soil properties in the short-term. We have also continued to develop WaderMap as a tool to collate information on different wader conservation activities and to display trends in wader population data.
- Biodiversity trends and changes across northern Europe are being driving by rising temperatures. A Europe-wide collaboration including SEFARI researchers analysed 161 biodiversity time series (over 15 to 91 years duration) covering 6,200 marine, terrestrial, and freshwater species from 21 European countries. The paper published in Nature Communications showed that local trends in biodiversity often deviate significantly from global patterns. In particular, the composition of species communities has undergone extensive changes at the local level. For northern Europe, the trend is towards an increase in diversity and species numbers which could be attributed to a combination of climate change (e.g., new species adapted to warmer conditions, spread of alien species and recovery from past disturbances and pollution (e.g., acidification). The study also showed declines in insect species abundances in the Atlantic bio-geo-region (UK, France, Belgium, Spain, Denmark and Germany), corroborating recent reports of worldwide declines of local terrestrial insect communities. The study emphasised the need to standardise biodiversity monitoring schemes and to integrate long-term biodiversity and environmental monitoring data, which will allow conservation measures to be better tailored at the local level.
- Enabling more sustainable food- and feed-systems: As part of an EU-wide team, a SEFARI researcher has discerned twenty-one enabling approaches to increase the production and consumption of home-grown legumes. These practical approaches present integrated strategies to benefit the environment, society, and the economy (Balázs et al., 2021). One such approach is processing beans for brewing and distilling, which demonstrates the environmental, nutritional, and commercial potential of the new, more sustainable agri-feed and -food systems (Black et al. 2021).
- Bere barley diversity contributes to sustainability and value chain diversification: SEFARI scientists have made further discoveries related to the importance of maintaining Bere barley diversity for its value to sustainability. Scotland’s traditional barley variety, Bere, has been shown to be particularly able to cope with micronutrient deficiency and traditional growth with seaweed as a fertiliser has driven the evolution of this ability (Brown et al. 2020). Genes associated with the ability of Bere to cope with manganese deficiency specifically have been identified (Cope et al. 2020). The importance of Bere as a heritage variety with value chain diversification credentials has been recognised in its inclusion in a successful bid for EU Horizon 2020 funding (RADIANT), which secured additional funding of €6M for the consortium.
- Biodiversity-function relationships in sustainable crop production: SEFARI scientists have produced a review article that links fundamental ecological understanding of the role of biodiversity in delivering ecosystem functions to the pressing need for developing sustainable crop production systems. The article reviews the potential benefits to multiple ecosystem functions and services from enhancing diversity in crop systems, for example through intercropping, and is relevant to research, as well as those interested in the development of sustainable farming policy.
Research has been performed to understand the mechanisms underpinning diversity benefits in arable plant communities including an experiment testing mixtures (up to six species) of cereals (barley, wheat), oil seeds (oil seed rape, linseed) and legumes (pea, faba bean) to understand how trait diversity contributes to improved productivity. A paper was published from the 2016 field trial that showed that increasing the number of barley genotypes in a mixture reduced the available niche space for weeds. The fate of newly reintroduced populations of Cicerbita alpina (alpine sow-thistle) has been followed and it appears that identity of the source population is an important determinant of survival. Material to re-establish the native fern Woodsia ilvensis (oblong woodsia) is ready for testing for drought-resistance. Fieldwork observations have been carried out at Glen Finglas to establish how long-term differences in grazing influence food quality for invertebrates and birds and the accompanying lab work is underway. Research on bere barley work has identified genetic markers that influence micronutrient uptake, revealing genetic diversity in traditional landraces that could benefit commercial barley by improving growth in nutrient-deficient soils. Modelling research has shown that maintenance of within-species diversity of insect herbivores (aphid pests) depends on the efficiency of their natural enemies. WaderMap, an application to bring together data on wader conservation projects and other environmental data, was successfully received and sampling in the large-scale liming experiment carried out. The two new bryophyte-based Ecosystem Health Indicators were launched on SEWeb accompanied by a SEFARI case study and the publication of the associated paper. Work on western oakwoods showed that long-term declines in lichen diversity have occurred over the last 50 years, whilst new work is focussed on identifying new sites for woodland regeneration to maximise potential colonisation by woodland specialists.
- Long-term consequences of upland grazing: SEFARI scientists have demonstrated that likely changes in grazing levels in the uplands (e.g. in response to changes in agricultural support payments, profitability of upland farming and conservation priorities) can take many years to affect upland plant communities. Specifically, it took 15 years for the tripling of sheep numbers or the complete removal of sheep to impact on most species. Some communities were very resistant to change, particularly those most attractive to grazers, which had lost plant species capable of responding to reduced grazing levels.
- Natural enemy regulation of diversity in insect herbivores. SEFARI scientists have shown evidence for a novel mechanism that could be responsible for maintaining genetic diversity within aphid populations. Aphids frequently shown resistance to parasitism by parasitoid wasps but the reason why susceptible and resistant aphids coexist has never been explained. Modelling work has shown that this could come about if parasitoid wasps suffer a fitness cost when switching between different aphid types. These findings are important for targeted biological control of aphid pests.
- Monitoring biodiversity through citizen science: SEFARI scientists have analysed citizen science data to document a five-decade decline in species that are associated with Scotland’s ancient woodlands. This was used as evidence to the State of Nature Report 2019 Solutions to reverse the trend were identified and have been used to inform the new strategy of the Atlantic Woodland Alliance consortium. This includes targeting efforts to the expansion of existing woodlands with old-growth structure, and which include populations of dispersal-limited species.
- Towards truly sustainable agri-food value-chains: RESAS-supported research on legumes (e.g. peas and beans) was recently reported in partnership with the EU TRUE project in the outcome of an EU-wide ‘consensus opinion of stakeholders’ exercise. Three recommended action areas were identified to help realise more-sustainable legume-supported agri-food systems around: investment in agri-food and -feed research and knowledge transfer; reducing reliance on the use of inorganic nitrogen fertiliser; plus greater foci on nutrition, health policies and public campaigns that promote the inclusion of home-grown legumes in the human diet.
- A toast to reduce Scotland’s protein import dependency: RESAS-supported work to help realise more cropping of pulses (e.g. peas and beans) in Scotland, carried out in partnership with Scottish-based SMEs, revealed that the use of home-grown peas to make gin also provided a high-protein ‘co-product’ for animal feed. The pea co-product can offset animal feed imports, and the two high-value products were achieved with a reduced carbon footprint (compared to conventional wheat gin). A commercial product (World’s first climate-positive gin) will be launched in February shortly after World Pulses Day (FAO-UN, Feb 10th).
Year 3 progress included a successful analysis of the barley:weed drought study and a new experiment on barley:weed interactions. The Cicerbita alpina (Alpine blue sow-thistle) reintroduction work made great strides in understanding the role of genetic diversity and restoration success, whilst the bere barley work has initiated the transfer of genes into elite varieties to cope with alkaline soils. A new fieldwork approach looking at management cascades through different trophic levels has started successfully at Glen Finglas and The work on Ecosystem Health Indicators has progressed well and is nearly ready for launching on SEWeb. Due to the cold spring, sowing of the grassland diversification experiment was delayed until May when conditions were more suitable. However, the dry summer meant that germinations was slow and consequently no monitoring was carried out in summer 2018. Research into effects of lime addition on trophic interactions continued to progress well, including the ongoing monitoring of three experimental study sites to contribute data on the effects of lime addition on soil characteristics, earth worm abundance, sward productivity, and vegetation diversity. Combined with the baseline data and ongoing monitoring this will provide data for later analyses to better understand the effects of lime addition on the availability of key soil invertebrates for breeding waders.
- Public engagement for biosecurity: Research on plant and animal biosecurity from the RBGE, James Hutton Institute, Moredun Research Institute, and SRUC was distilled into an interactive, permanent public exhibit at the John Hope Gateway. SEFARI work is presented in a series of interactive panels demonstrating how plant and animal diseases move around the world, and actions the public can take to protect Scotland’s natural and economic assets. This exhibit is prominently placed in the entrance to the RBGE (694,000 international and local visitors in 2016), with portable panels in construction to further spread the message at public events across Scotland.
- Major new database on multi-scale plant diversity published: The James Hutton Institute was the main UK contributor to a new international database of vegetation data - GrassPlot. The difference between this database and others is that all the plot data were collected at multiple scales. The main aim of collecting and publishing these data is to facilitate studies on the scale- and taxon-dependency of biodiversity patterns and drivers along macroecological gradients. For instance, it will allow researchers to analyse how scale dependent relationships such as the species-area curve vary with climate.
- Importance of crop mixtures research brought to the fore through Parliamentary Reception and new publication: Research on potential benefits of crop mixtures for sustainable food production was highlighted during a SEFARI Showcase event at the Scottish Parliament. The event’s keynote address was given by the Cabinet Secretary for Rural Economy and Connectivity, Fergus Ewing MSP. In addition, concepts developed through WP1.3 work on crop mixtures contributed to delivery of a new research paper in Nature Ecology and Evolution. This shows that some of the effects which might help to deliver benefits from crop mixtures evolve best in species rich plant communities, indicating benefits for sustainability from conserving species.
- Evolving for mixture benefits: Concepts developed through our crop mixtures work (RD1.3.1 and RD2.3.8) concerning the beneficial interactions between neighbouring species have contributed to delivery of a new research paper by Schöb et al. This study shows for the first time that some of the effects which help to deliver benefits from crop mixtures evolve best in a species rich plant community. Importantly this work demonstrates how higher diversity helps promote ecosystem functions, and that breeding for crop mixtures (a potential sustainable production approach) may need novel trait combinations which only emerge in species-rich systems.
- Establishing landraces as living heritage: SEFARI scientists in collaboration with Archaeologists from University of Sheffield have identified and confirmed the origins of barley landraces in the archaeological record using morphometrics and genetics, helping to establish its status as living heritage and helping secure its commercial future through providing the grain with heritage credentials that can be used in marketing of products.
- Ancient barley landraces adapted to marginal soils demonstrate exceptional tolerance to micronutrient limitation: SEFARI scientists in collaboration with University of Copenhagen and University of Highlands and Islands, Orkney have identified a unique Mn efficient phenotype in Bere landraces that can be potentially used to cope with micronutrient deficiencies seen in crops grown in marginal calcareous soils in the western and northern isles of Scotland and in many of the cereal producing lands on a global scale.
Year 2 progress included an experiment to investigate how increasing the genetic diversity of barley improved the tolerance of the crop to drought; this was not the case but increasing diversity suppressed weeds and reduced disease incidence. Alpine blue sow-thistle Cicerbita alpina was re-introduced into three new sites in the Cairngorms. The translocated populations include an experimental approach to test the resilience of different genotypes with the aim of increasing in situ seed-set (see Highlights below). Baseline data of the effects of lime addition to trophic interactions, in particular its impacts on potential invertebrate prey of wading birds, have been collected. Information on grass productivity and quality is also being collected. We have developed additional Ecosystem Health Indicators relating to nitrogen pollution and climate change based on moss and liverwort records in the National Biodiversity Network and their habitat preferences and provided initial results that will feed into the Scottish Biodiversity Strategy. Based on information on epiphyte species distributions, a report identifying priority areas for restoration has been used to inform the development of the Atlantic Woodland Alliance.
- Potential for crop improvement in marginal soils: New research has shown that extant barley landraces selected over many generations on marginal soils have adapted to tolerate soils with limited Manganese (Mn) availability. By contrast, modern elite varieties on similar soils fail to complete their lifecycle, due to their poor Mn efficiency. Landraces derived from regions with reduced soil fertility constitute a valuable resource of untapped adaptive genetic variation, with an opportunity to identify the key adaptive traits to underpin crop improvement in marginal soils.
- Rare plants may be helped by crops: A study investigating the interactions between a rare vascular plant - Valerianella rimosa - (broad fruited corn salad) and a barley crop has shown that the conditions found within the crop might help promote the establishment of the rare plant at the start of the growing season. When combined with the negative impact of the crop on common weeds, these findings indicate that crops may play a role in the conservation of some rare vascular plants by creating space in the farmland weed community.
- Reintroduction of a rare plant to the Cairngorms: Studies of genetic diversity in populations of the rare Alpine blue sow-thistle have enabled the reintroduction of the species at three new locations in the Cairngorms. Contributing to both the Cairngorms Nature Action Plan and Priority Project 9 of the 2020 Route Map for Scotland’s Biodiversity, the translocated populations included an experimental approach to test the resilience of different genotypes with the aim of increasing the viable seed-set.
- The destructive effects of Rhododendrons: Rhododendron ponticum shades out native understorey flora and has been the focus of removal by conservation groups. A paper on its effects in Celtic rainforests showed that even after removal of Rhododendron, woodlands did not automatically revert to their pre-invaded state; reseeding with native flowers and grasses is required to fully restore them. Media publicity included interviews for Radio Wales, Radio Scotland and Radio 4 (World at One) and reports in the Daily Mail, The Times, Horticulture Week, Sunday Post, Press and Journal, the Scotsman and others.
Progress in the Research Deliverable includes successful experiments on diversity-driven trophic interactions; these showed a postive relationship between weed species richness and barley productivity and that increasing barley diversity reduced weed performance through increased competition for light. Trials in Orkney and Dundee have been used to identify differnces in bere barley performance and potential useful traits to breed into modern barley cultivars. A long-term experiment assessing ecosystem effects of liming has been set up, with a particular focus on assessing the impact of liming on the provision of prey to wading birds, the improvement of quality and productivity of the grasslands and the impact of lime on soil carbon. Plant material to re-introduce Cicerbita alpina has been bred from extant Scottish and Norwegian populations; with propogation carried out to ensure no potential for disease spread. Analysis of a long-term experiment on trophic interactions in the uplands has revealed the importance of management on both insectivourous bird:invertebrate prey and predator:vole interactions. Collaboration with the Scottish Biodiversity Strategy indicators group has led to further refinement of the Natural Capital Asset Index. Connectivity work has produced indices of connectivity for woodlands in Scotland, used to start analyses of priority restoration sites for Atlantic Oak woodlands.
- A new paper in Plant Ecology & Diversity demonstrates how arable weed diversity has negative impacts on new plant species establishing (especially rare plant species).
- Workshop was held to improve collaboration between SRP research and major Scottish environmental indicators initiatives, including Ecosystem Health Indicators, Natural Capital Asset Index and the Soil Monitoring Action Plan; the outcome is an improved alignment of SRP research to end-user needs and which will support the future development of these initiatives.
- WP1.3 researchers helped deliver the Scottish Biodiversity Strategy Science Conference at RBGE in November. Over 100 delegates attended, including government, agencies, eNGOs and the public (further information here).
- The Ecosystems and Land Use Stakeholder Engagement Group (WP1.3 & 1.4) had its first meeting at Victoria Quay in November. Fifty-five stakeholders from government, agencies, Non-Governmental Organisations, membership organisations and universities attended and were engaged with ongoing work and identifying cooperation opportunities.
The final year of this programme will partly see effort to catch up with Covid-19 effected work but also extension of current work into new areas.
The work on crop species mixtures will be extended through a pilot study for participatory research to prototype a ‘Living Lab’ for Citizen Science in Agroecology (see also RD2.3.8). It will include data from previous trials and be supplemented with farmer collected data from on-farm trials in 2021. Data will be published using an open science platform. Population rescue work for Cicerbita alpina is being extended to cover Woodsia ilvensis, and research on improving woodland diversity will include the development of a toolkit for assessing forest biodiversity and woodland structure. The work on Bere barley will extend to testing their resistance against a wider range of environmental stresses and the work across the life of the programme publicised through a podcast or video. The upland grazing work at Glen Finglas will take a new direction in linking earth observation data to the ground-based data we have been collected so that we have a better measure of site productivity to use in our analyses. Additional upland focussed work will extend the “Phase 2 Grouse Research” work on assessing the area of heather burning in Scotland to cover all of the relevant parts of Scotland. The work on sward diversification missed monitoring last year, so monitoring of establishment and spread of introduced species will continue, but it will be supplemented by a review of mob stocking/tall-grass grazing.
- Brooker et al. (2017) Crop presence, but not genetic diversity, impacts on the rare arable plant Valerianella rimosa. Plant Ecology & Diversity
- Popular articles on the role of diversity in cropping systems, restoration after rhododendron control and the strengths of heritage barley have been published in the Scottish Consortium for Rural Research Newsletter, Issue 89, Winter 2017.
- Mládková, P., Mládek, J., Hejduk, S., Hejcman, M. &Pakeman, R.J. (2018) Calcium plus magnesium indicates digestibility: the significance of the second major axis of plant chemical variation for ecological processes. Ecology Letters.
- Liming trial and predator monitoring work were presented at the "The Challenge of Marginal Land: efficient utilisation of the managed natural environment" workshop, Glensaugh, 13 Sept 2017.
- Schöb et al. (2018) Evolution of facilitation requires diverse communities (Nature Ecology and Evolution).
- Brooker, R.W., Hewison, R., Mitchell, C., Newton, A.C., Pakeman, R.J., Schöb, C., Karley, A.J. (2018) The role of crop genetic diversity in determining plant community resilience to experimental drought.
- A film on Cicerbita alpina has been produced and is currently shown (June 2018 – open ended) at the John Hope Gateway, Royal Botanic Garden Edinburgh.
- Williams, L. & Ellis, C.J. (2018) Ecological constraints to 'old-growth' lichen indicators: niche specialisation or dispersal limitation? Fungal Ecology, 34: 20-27.
- Ellis, C.J. (2017) When is translocation required for the population recovery of old-growth epiphytes in a reforested landscape? Restoration Ecology, 25: 922-932.
- Eaton at al. (2018) Adding small species to the big picture: species distribution modelling in an age of landscape scale conservation. Biological Conservation, 217: 251-258.
- Eaton et al. (2018) A method for the direct detection of airborne dispersal in lichens. Molecular Ecology Resources, 18: 240-250.
- Ellis, C.J. & Eaton, S. (2018) The biogeography of climate change risk for Scotland’s woodland biodiversity: epiphytes. Scottish Geographical Journal, 134: 257-267.
- Pakeman, R.J.; Brooker, R.W.; Karley, A.J.; Newton, A.C.; Mitchell, C.; Hewison, R.L.; Pollenus, J.; Guy, D.C.; Schöb, C. (2020) Increased crop diversity reduces the functional space available for weeds. Weed Research 60: 121-131.
- Pakeman, R.J., Brooker, R.W., O'Brien, D. and Genney, D. (2019) Using species records and ecological attributes of bryophytes to develop an ecosystem health indicator. Ecological Indicators, 104, 127-136. The bryophyte indicator is now live on the SE Web platform: https://www.environment.gov.scot/
- Donnelly, D.; Fielding, D.; Newey, S. (2019) WaderMap: Collaboration and Cooperation in Action(Version 1.3).
- Ellis, C.J. & Coppins, B.J. (2019) Five decades of decline for old-growth indicator lichens in Scotland. Edinburgh Journal of Botany, 76, 319-331.
- Preedy, K.F., Chaplain, M.A., Leybourne, D.J., Marion, G. and Karley, A.J. (2020) Learning‐induced switching costs in a parasitoid can maintain diversity of host aphid phenotypes although biocontrol is destabilized under abiotic stress. Journal of Animal Ecology, 89, 1216-1229.
- Temesgen, D., Maluk, M., James, E. K., Iannetta, P. P. M., Assefa, F. 2019) The functional characterisation of soybean (Glycine max L.) rhizospheric bacteria indigenous to Ethiopian soils. African Journal of Agricultural Research 14, 1659-1673.
- Leybourne, D., Valentine T.A., Robertson, J.A.H., Pérez-Fernández, E., Main, A.M., Karley, A.J. & Bos, J.I.B. (2019) Defence gene expression and phloem quality contribute to mesophyll and phloem resistance against aphids in wild barley. Journal of Experimental Botany 70 (15), 4011-4026. doi: 10.1093/jxb/erz163
- Schmidt, S.B., Hedley, P.E., George, T.S., Brown, L.K., Booth, A., Wishart, J., Martin, P., Russell, J. & Husted, S. (2019) Ancient barley landraces adapted to marginal soils demonstrate exceptional tolerance to manganese limitation. Annals of Botany, 123: 831–843.
- Wallace, M., Bonhomme, V., Russell, J. et al. (2019) Searching for the origins of bere barley: a geometric morphometric approach to cereal landrace recognition in archaeology. Journal of Archaeological Method Theory, 26: 1125-1142
- Brooker, R. et al. (2021) Facilitation and biodiversity–ecosystem function relationships in crop production systems and their role in sustainable farming. Journal of Ecology
- Losapio, G. et al. (2021) Network motifs involving both competition and facilitation predict biodiversity in alpine plant communities. Proceedings of the National Academy of Sciences of the USA (PNAS), 118, Article No. e2005759118.
- Vasconcelos, M., Gomes A., Pinto, E., Ferreira, H., Vieira, E., Pimenta, A., Santos, C.S., Balázs, B., Kelemen, E., Hamann, K., Williams, M., Iannetta, P.P.M. (2020) The push-, pull- and enabling – capacities necessary for legume grain inclusion into sustainable agri-food systems and healthy diets. Chapter 1, in ‘Hidden hunger and the transformation of food systems. How to combat the double burden of malnutrition?’ Biesalski HK (ed). World Review of Nutrition and Dietetics (Basel, Karger), 121.
- Karavidas, I., Ntatsi, G., Ntanasi, T., Vlachos, I., Tampakaki A., Iannetta, P., Savvas, D. (2020) Comparative assessment of different crop rotation schemes for organic common bean production. Agronomy 10, 1269.
- Malm, L.E., Pearce‐Higgins, J.W., Littlewood, N.A., Karley, A.J., Karaszewska, E., Jaques, R., Pakeman, R.J., Redpath, S.M., Evans, D.M. (2020) Livestock grazing impacts components of the breeding productivity of a common upland insectivorous passerine: Results from a long‐term experiment. Journal of Applied Ecology, 57, 1514-1523.
- Pakeman, R.J. The Glen Finglas Experiment and the slow pace of change in upland communities.
- Pakeman, R.J.; Fielding, D.A. A functional assessment of the impact of changing grazing management of upland grassland mosaics. Applied Vegetation Science 23, 539-550.
- Pakeman, R.J. Plants, invertebrates and birds in upland grassland: effects of grazing management. Biodiversity in Grassland Webinar. British Society of Animal Science, British Grassland Society and the Stapledon Trust, 26 January 2021."
- Fielding, D., Newey, S, & Pakeman, R. (2021) Assessing the impacts of lime application to pastures on soils, biodiversity, and forage quality.
- Donnelly, Fielding and Newey (2021) WaderMap: Collaboration and cooperation in action (Version 1.7).