Many aspects of plant development are regulated by antagonistic interactions between the plant hormones auxin and cytokinin, but the molecular mechanisms of this interaction are not understood. To test whether cytokinin controls plant development through inhibiting an early step in the auxin response pathway, we compared the effects of cytokinin with those of the dgt (diageotropica) mutation, which is known to block rapid auxin reactions of tomato (Lycopersicon esculentum) hypocotyls. Long-term cytokinin treatment of wild-type seedlings phenocopied morphological traits of dgt plants such as stunting of root and shoot growth, reduced elongation of internodes, reduced apical dominance, and reduced leaf size and complexity. Cytokinin treatment also inhibited rapid auxin responses in hypocotyl segments: auxin-stimulated elongation, H ϩ secretion, and ethylene synthesis were all inhibited by cytokinin in wild-type hypocotyl segments, and thus mimicked the impaired auxin responsiveness found in dgt hypocotyls. However, cytokinin failed to inhibit auxin-induced LeSAUR gene expression, an auxin response that is affected by the dgt mutation. In addition, cytokinin treatment inhibited the auxin induction of only one of two 1-aminocyclopropane-1-carboxylic acid synthase genes that exhibited impaired auxin inducibility in dgt hypocotyls. Thus, cytokinin inhibited a subset of the auxin responses impaired in dgt hypocotyls, suggesting that cytokinin blocks at least one branch of the DGT-dependent auxin response pathway.The balance between auxin and cytokinin controls a wide range of processes in plant development, including the formation of roots, shoots, and callus tissue in vitro (Skoog and Miller, 1957), the outgrowth of shoot axillary buds (Sachs and Thimann, 1967), and the formation of lateral roots (Wightman et al., 1980; Hinchee and Rost, 1986). Mutual control of active auxin and cytokinin pools, interactive control of gene expression, and posttranslational effects have been described as possible mechanisms underlying such physiological interactions (Coenen and Lomax, 1997). However, the relationship between classical hormone interactions at the physiological level and molecular auxin-cytokinin interactions is presently not well defined.
Auxin-Cytokinin Interactions during Hypocotyl ElongationAuxin-cytokinin interactions can be observed in the elongation response of dicot hypocotyl segments. Auxin-induced elongation of sunflower (Helianthus annuus; DeRopp, 1956) and soybean (Glycine max; Vanderhoef et al., 1973; Vanderhoef and Stahl, 1975) hypocotyl segments is inhibited in the presence of cytokinins. This inhibition is detectable within 10 min of cytokinin addition and is not mediated by changes in ethylene synthesis (Vanderhoef and Stahl, 1975). Based on the time course of auxin-induced elongation growth in the presence and absence of cytokinin, Vanderhoef and Stahl (1975) proposed that cytokinin selectively inhibits the later phase of auxininduced elongation and may not influence rapid growth processes mediate...