'Haying-off', the negative grain yield response of dryland wheat to nitrogen fertiliser. I. Biomass, grain yield, and water use

'Haying-off' was studied by comparing wheat responses to applied nitrogen (N) at 3 sites in southern New South Wales, which differed in the amount and timing of rainfall during crop growth. At a site where the crops encountered little water deficit, dry grain yield increased from 607 g/m² for a low-N control crop to 798 g/m² for a high-N crop. At a site with severe terminal drought, dry grain yield decreased 24% from 374 g/m² for the control, to 284 g/m² for the highest N crop. At the third site, yields increased with small applications of N, whereas greater applications resulted in a negative yield response. At the 2 latter sites, the crops that showed decreased yield with applied N had clearly hayed-off. At all sites, irrespective of water status, N application resulted in increases in biomass at anthesis, spike density, kernels per spike, and kernel number. Kernel weight decreased in response to additional N at all sites, but most markedly at the haying-off sites, where it decreased by up to 38%. Harvest index increased in response to N at the high-rainfall site, but decreased in crops that hayed-off. Grain protein increased in response to N at all sites, with a range from 9% to 18% at the haying-off sites. The apparent retranslocation of assimilates to grain contributed 37-39% of grain yield (depending on N supply) at the high-rainfall site, compared with 75-100% at the haying-off sites. In contrast, when apparent retranslocation was expressed in relation to biomass at anthesis, it remained relatively constant, amounting to 23-26% at the high-rainfall site and 24-28% when crops hayed-off. By anthesis, high-N crops extracted more soil water than the low-N crops. By maturity the most severely hayed-off crop had extracted 10 mm less soil water than a low-N crop, but at the high rainfall site the high-N crops extracted 20 mm more soil water than the control crops. The weather conditions between anthesis and physiological maturity were relatively mild, with no daily maximum temperatures above 30°C and no sudden increases in evaporative demand. Thus, there appeared to be 3 processes leading to haying-off. Firstly, the results confirm previous studies showing that haying-off was associated with reduced post-anthesis assimilation in response to a lack of soil water. The water deficit was due to vigorous vegetative growth stimulated by a high level of soil N and was not associated with heat shocks or sudden increases in evaporation. Secondly, the most severely hayed-off crop failed to extract soil water fully, leading to a further reduction in post-anthesis assimilation. Thirdly, there was inadequate apparent retranslocation of pre-anthesis reserves to compensate for the lack of post-anthesis assimilation.