RESUMEN

The role of hormones in mediating tropic responses has been a central question in plant biology. Another key issue concerns how interactions between hormones regulate plant responses. In the September 2007 issue of Physiologia Plantarum, we published a paper relevant to both these questions.1 This paper focuses on gravitropism in the barley leaf sheath pulvinus. The results support the Cholodny-Went theory on hormones and tropic responses, and highlight how an environmental factor (gravity) appears to first affect auxin content and consequently that of bioactive gibberellins (GAs). It appears that while GAs do not actually trigger the gravitropic bending of barley pulvini, they do act to magnify the bending response.

RESUMEN

The auxin indole-3-acetic acid (IAA) is known to promote the biosynthesis of active gibberellins (GAs) in barley (Hordeum vulgare). We therefore investigated the possibility that this interaction might contribute to the gravitropic response of barley leaf sheath pulvini. Barley plants at the inflorescence stage were gravistimulated for varying times, and the pulvini were then separated into upper and lower halves for quantification of IAA and GAs by GC-MS. Consistent with the Cholodny-Went theory, the lower portion contained more IAA than did the upper portion. This difference was detected as early as 2.5 h after the start of gravistimulation, and bending was also observed at this stage. At later time points tested (6 h and 24 h), but not at 2.5 h or 3 h, the higher auxin content of the lower half was associated with a higher level of GA(1), the main bioactive GA in barley. Consistent with that result, the expression of Hv3ox2, which encodes a key enzyme for the conversion of GA(20) to GA(1), was higher in the lower side than in the upper, after 6 h. It is suggested that in gravistimulated leaf sheath pulvini, auxin accumulates in the lower side, leading to a higher level of GA(1), which contributes to the bending response. Further evidence that GAs play a role in the gravitropic response was obtained from GA-related mutants, including the elongated sln1c mutant, in which GA signalling is constitutive. Pulvinar bending in the sln1c mutant was greater than in the wild-type. This result indicates that in the lower side of the gravistimulated pulvinus, the relatively high level of bioactive GA facilitates, but does not mediate, the bending response.

Asunto(s)

RESUMEN

Multiple gibberellins (GAs) were quantified in the stems of intact, decapitated, and decapitated auxin-treated barley (Hordeum vulgare) plants. Removal of the developing inflorescence reduced the endogenous levels of indole-3-acetic acid (IAA), GA(1), and GA(3) and increased the level of GA(29) in internodal and nodal tissues below the site of excision. Application of IAA to the excised stump restored GA levels to normal in almost all cases. The conversion of [(14)C]GA(20) to bioactive [(14)C]GA(1) and of [(14)C]GA(5) to bioactive [(14)C]GA(3) was reduced by decapitation, and IAA application was able to restore conversion rates back to the levels found in intact plants. The amount of mRNA for the principal vegetative 3-oxidase (converting GA(20) to GA(1), and GA(5) to GA(3)) was decreased in decapitated plants and restored by IAA application. The results indicate that the inflorescence of barley is a source of IAA that is transported basipetally into the internodes and nodes where bioactive GA(1) and GA(3) are biosynthesized. Thus, IAA is required for normal GA biosynthesis in stems, acting at multiple steps in the latter part of the pathway.

Asunto(s)

RESUMEN

To investigate gibberellin (GA) biosynthesis in mature tissue of pea (Pisum sativum L.) in the absence of potentially GA-producing meristematic tissue we grafted wild-type scions to rootstocks of the GA-deficient ls-1 mutant and later decapitated the shoot. After 2 d, decapitated shoots contained as much GA19 (a precursor of the bioactive GA1) as comparable tissue from intact plants, even though applied [14C]GA19 was metabolised rapidly during this time. These results show that the pool size of endogenous GA19 was maintained, probably by de novo GA19 synthesis. We also found that the LS gene, which catalyses an early step in GA biosynthesis, is expressed in mature tissue, as are the shoot-expressed GA 20-oxidase and GA 3-oxidase genes. Nevertheless, mature tissue contained very low levels of GA1 and GA20 compared with immature tissue. Levels of GA19, GA29 and GA8 were less affected by tissue age. Metabolism studies using 14C-labelled GAs indicated that mature tissue rapidly converted GA19 to GA20 and GA20 to GA1; the latter step was promoted by IAA. However, the 2-oxidation steps GA1 to GA8, GA20 to GA29 and GA29 to GA29-catabolite appear to proceed very rapidly in mature tissue (regardless of IAA content), and we suggest this is the reason why GA1 and GA20 do not accumulate. This is supported by the high level of expression of a key GA 2-oxidase gene in mature tissue.

RESUMEN

Recently it was discovered that auxin promotes gibberellin (GA) biosynthesis in decapitated stems of pea (Pisum sativum L.) and tobacco (Nicotiana tabacum L.), and here we review the evidence for this interaction. We also discuss the possible relationship between auxin and the mechanisms by which bioactive GAs (such as GA1) regulate their own levels, and the implications of the auxin-GA interaction for the control of plant growth. It is now possible to envisage auxin as a messenger linking the apical bud with the biosynthesis of active GAs in the expanding internodes. Finally, new evidence is presented that the promotion of growth by GA1 does not depend on GA1-induced increases in auxin content.