Globalisation brings pressure on regional production systems to become regional production and innovation systems in order to maintain their competitiveness. However, the heterogeneity of the companies in these systems makes it difficult to respond to these challenges and obliges the response to be collective and mesoeconomic. To date, there has been a defence of the leadership of some private companies and other intermediaries such as business associations as vehicles of this process, but the part that cooperatives play in it has barely been studied. This paper therefore pursues a double aim. Firstly, it undertakes a novel conceptualisation of the cooperatives? capacity for transforming regional production systems into innovation systems. Combining an evolutionary approach with that of governance and the collective entrepreneurship of the social economy brings to light a truly cooperative ?third way? (between private and public) to transform these systems through a cooperative model of an innovation system that is capable of generating new meso-rules. Secondly, based on 218 in-depth interviews with different players in the Valencian citrus industry, it shows how the second-tier cooperative ANECOOP has led the shaping of a cooperative model of an innovation system. As a result, in response to the new demands of growers, packers and marketers for scientific and technical knowledge, new routines and forms of organisation have arisen through the interaction and redefinition of the communities of practice and the epistemic communities.