Soil conditions and plant growth'

Plants can respond to soil conditions in ways that can not readily be explained in terms of the ability of the roots to take up water and nutrients. Roots may sense difficult conditions in the soil and thence send inhibitory signals to the shoots which harden the plants against the consequences of a deteriorating or restrictive environment, especially if the plants' water supply is at risk. Generally, this behaviour can be interpreted as feedforward responses to the soil becoming too dry or too hard, or to the available soil volume being very small as with bonsai plants, or to roots' becoming infected with pathogens. However, soil that is too soft or in which the roots are forced to grow in very large pores can also induce large conservative responses, the significance of which is unclear. The inhibitory signals may affect stomatal conductance, cell expansion, cell division and the rate of leaf appearance. Their nature is still under debate, and the debate is becoming increasingly complex, which probably signifies that a network of hormonal and other responses is involved in attuning the growth and development of a plant to its environment.