Opportunities for improving intrinsic water use efficiency in C₄ plants under climate change

C₄ photosynthesis is inherently efficient, saturating at low intercellular CO₂ (C_i) and operating under low stomatal conductance (g_s), resulting in high intrinsic water use efficiency (iWUE = assimilation rates, (A)/ stomatal conductance (g_s)). While iWUE is generally higher in C₄ than in C₃ plants, future climate scenarios, marked by increasing atmospheric CO₂, temperatures, and aridity, present ongoing pressures on crop productivity in C₄-dominated environments. This review explores the physiological, anatomical, and environmental factors controlling iWUE in C₄ plants, with a focus on balancing productivity and resilience under climate change. Using process models of photosynthesis and stomatal behaviour, we assess how iWUE varies with photosynthetic limitations, highlight the central role of stomata in controlling iWUE, and predict the physiological adjustments needed to optimize carbon gain per unit water loss. Recent evidence points to leaf width as a useful trait influencing leaf energy balance, temperature, and transpiration. We propose integrated physiological and breeding strategies to optimize crop iWUE and yield under climate stress, supported by emerging sensing and selection technologies.