For most plants survival depends upon the capacity of root tips to sense and move towards water and other nutrients in the soil. Because land plants cannot escape environmental stress they use developmental solutions to remodel themselves in order to better adapt to the new conditions. The primary site for perception of underground signals is the root cap (RC). Plant roots have positive hydrotropic response and modify their growth direction in search of water. Using a screening system with a water potential gradient, we isolated ano hydrotropic response (nhr) semi-dominant mutant of Arabidopsis that continued to grow downwardly into the medium with the lowest water potential contrary to the positive hydrotropic and negative gravitropic response seen in wild type-roots. The lack of hydrotropic response of nhr1roots was confirmed in a system with a gradient in air moisture. The root gravitropic response of nhr1 seedlings was significantly faster in comparison with those of wild type. The frequency of the waving pattern in nhr1 roots was increased compared to those of wild type. nhr1 seedlings had abnormal root cap morphogenesis and reduced root growth sensitivity to abscisic acid (ABA) and the polar auxin transport inhibitor N-(1-naphtyl)phtalamic acid (NPA). These results showed that hydrotropism is amenable to genetic analysis and that an ABA signaling pathway participates in sensing water potential gradients through the root cap.

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While water shortage remains the single-most important factor influencing world agriculture, there are very few studies on how plants grow in response to water potential, i.e., hydrotropism. Terrestrial plant roots dwell in the soil, and their ability to grow and explore underground requires many sensors for stimuli such as gravity, humidity gradients, light, mechanical stimulations, temperature, and oxygen. To date, extremely limited information is available on the components of such sensors; however, all of these stimuli are sensed in the root cap. Directional growth of roots is controlled by gravity, which is fixed in direction and intensity. However, other environmental factors, such as water potential gradients, which fluctuate in time, space, direction, and intensity, can act as a signal for modifying the direction of root growth accordingly. Hydrotropism may help roots to obtain water from the soil and at the same time may participate in the establishment of the root system. Current genetic analysis of hydrotropism in Arabidopsis has offered new players, mainly AHR1, NHR1, MIZ1, and MIZ2, which seem to modulate how root caps sense and choose to respond hydrotropically as opposed to other tropic responses. Here we review the mechanism(s) by which these genes and the plant hormones abscisic acid and cytokinins coordinate hydrotropism to counteract the tropic responses to gravitational field, light or touch stimuli. The biological consequence of hydrotropism is also discussed in relation to water stress avoidance.

Roots are highly plastic and can acclimate to heterogeneous and stressful conditions. However, there is little knowledge of the effect of moisture gradients on the mechanisms controlling root growth orientation and branching, and how this mechanism may help plants to avoid drought responses. The aim of this study was to isolate mutants of Arabidopsis thaliana with altered hydrotropic responses. Here, altered hydrotropic response 1 ( ahr1 ), a semi-dominant allele segregating as a single gene mutation, was characterized. ahr1 directed the growth of its primary root towards the source of higher water availability and developed an extensive root system over time. This phenotype was intensified in the presence of abscisic acid and was not observed if ahr1 seedlings were grown in a water stress medium without a water potential gradient. In normal growth conditions, primary root growth and root branching of ahr1 were indistinguishable from those of the wild type (wt). The altered hydrotropic growth of ahr1 roots was confirmed when the water-rich source was placed at an angle of 45° from the gravity vector. In this system, roots of ahr1 seedlings grew downward and did not display hydrotropism; however, in the presence of cytokinins, they exhibited hydrotropism like those of the wt, indicating that cytokinins play a critical role in root hydrotropism. The ahr1 mutant represents a valuable genetic resource for the study of the effects of cytokinins in the differential growth of hydrotropism and control of lateral root formation during the hydrotropic response.

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