Review: Exploring correctness, usefulness, and feasibility of potential physiological operational welfare indicators for farmed insects to establish research priorities.

While insects are already the largest group of terrestrial food and feed livestock animals in terms of absolute number of individuals, the insect farming industry is expected to continue growing rapidly in order to meet the nutritional demands of the human population during the 21st century. Accordingly, consumers, producers, legislators, and industry-adjacent researchers have expressed interest in further research and assessment of farmed insect welfare. Operational indicators of animal welfare are those that can be used to putatively assess the welfare of animals in the absence of true indicators of affective state (e.g., valenced/emotional state) and are commonly used for farmed vertebrate livestock species; however, significant behavioral and physiological differences between vertebrates and insects means these indicators must be examined for their correctness, usefulness, and feasibility prior to use with insect livestock. The most valuable operational welfare indicators would (1) correctly correspond to the insect's putative welfare state; (2) provide useful information about what is affecting the insect's welfare; and (3) be feasible for deployment at a large scale on farms. As there are many possible indicators that could be further researched in insects, evaluating the likely correctness, feasibility, and usefulness of these indicators in insects will allow researchers to prioritize which indicators to investigate first for use on farms. Thus, in this review, we explore whether physiological or somatic indicators of farmed vertebrate welfare, including whole-body, immune, neurobiological, and respiratory/cardiac indicators, may be correct, feasible, and useful for assessing farmed insect welfare. We review insect physiological systems, as well as any existing, welfare-relevant data from farmed or closely related insects. We end by proposing a priority list for physiological, operational welfare indicators that are most likely to correctly, usefully, and feasibly assess farmed insect welfare, which may guide indicator validation research priorities for insect welfare scientists.