TRANSPORTER 2 (SUT2), SUCROSE TRANSPORTER 4 (SUT4/SUC4) 27 Abstract (max 250 words): 28The circadian clock is considered a key target for crop improvement because it controls 29 metabolism and growth in Arabidopsis. Here, we show that the clock gene EARLY 30 FLOWERING 3 (ELF3) controls vegetative growth in Arabidopsis but not in the cereal crop 31 barley. Growth in Arabidopsis is determined by the degradation of leaf starch reserves at 32 night, which is controlled by ELF3. The vegetative growth of barley, however, is determined 33 by the depletion of leaf sucrose stores through an exponential kinetics, presumably catalyzed 34 by the vacuolar sucrose exporter SUCROSE TRANSPORTER 2 (SUT2). This process depends 35 on the sucrose content and the nighttime temperature but not on ELF3. The regulation of 36 starch degradation and sucrose depletion in barley ensures efficient growth at favorable 37 temperature as stores become exhausted at dawn. On cool nights, however, only the starch 38 degradation rate is compensated against low nighttime temperatures, whereas the sucrose 39 depletion rate is reduced. This coincides with reduced biomass in barley but not in 40Arabidopsis after growth in consecutive cool nights. The sucrose depletion metabolism 41 determines growth in the cereal crops barley, wheat, and rice but is not generally conserved 42 in monocot species and is not a domestication-related trait. Therefore, the control of growth 43 by endogenous (clock) versus external factors (temperature) is species-specific and depends 44 on the predominant carbohydrate store. Our results give new insights into the physiology of 45 growth in cereals and provide a basis for studying the genetics and evolution of different 46 carbohydrate stores and their contribution to plant productivity and adaptation. 47Significance Statement (120 words max): 48The circadian clock controls growth in the model plant Arabidopsis thaliana by regulating the 49 starch degradation rate so that reserves last until dawn. This prevents nocturnal starvation 50 until photosynthesis resumes. The cereal crops barley, wheat and rice, however, 51 predominantly consume sucrose instead of starch as carbohydrate source. We find that 52 carbohydrate supply from sucrose at night is regulated by enzyme kinetics and night-time 53 temperature, but not the circadian clock. We postulate that the regulation of growth depends 54 on the predominant carbohydrate store, where starch degradation is controlled by 55 endogenous cues (clock) and sucrose depletion by external factors (temperature). These 56 differences in the regulation of carbohydrate availability at night may have important 57 implications for adapting crops yields to climate change. 58 \body 59Introduction 60