Members of the homeodomain leucine zipper (HDZip) family of transcription factors are present in a wide range of plants, from mosses to higher plants, but not in other eukaryotes. The HDZip genes act in developmental processes, including vascular tissue and trichome development, and several of them have been suggested to be involved in the mediation of external signals to regulate plant growth. The Arabidopsis (Arabidopsis thaliana) genome contains 47 HDZip genes, which, based on sequence criteria, have been grouped into four different classes: HDZip I to IV. In this article, we present an overview of the class I HDZip genes in Arabidopsis. We describe their expression patterns, transcriptional regulation properties, duplication history, and phylogeny. The phylogeny of HDZip class I genes is supported by data on the duplication history of the genes, as well as the intron/exon patterning of the HDZip-encoding motifs. The HDZip class I genes were found to be widely expressed and partly to have overlapping expression patterns at the organ level. Further, abscisic acid or water deficit treatments and different light conditions affected the transcript levels of a majority of the HDZip I genes. Within the gene family, our data show examples of closely related HDZip genes with similarities in the function of the gene product, but a divergence in expression pattern. In addition, six HDZip class I proteins tested were found to be activators of gene expression. In conclusion, several HDZip I genes appear to regulate similar cellular processes, although in different organs or tissues and in response to different environmental signals.

The homeodomain leucine zipper (HD-Zip) genes encode transcription factors that have diverse functions in plant development and have often been implicated in stress adaptation. The HD-Zip genes are the most abundant group of homeobox (HB) genes in plants and do not occur in other eukaryotes. This paper describes the complete annotation of the HD-Zip families I, II and III from rice and compares these gene families with Arabidopsis in a phylogeny reconstruction. Orthologous pairs of rice and Arabidopsis HD-Zip genes were predicted based on neighbour joining and maximum parsimony (MP) trees with support of conserved intron-exon organization. Additionally, a number of HD-Zip genes appeared to be unique to rice. Searching of EST and cDNA databases and expression analysis using RT-PCR showed that 30 out of 31 predicted rice HD-Zip genes are expressed. Most HD-Zip genes were broadly expressed in mature plants and seedlings, but others showed more organ specific patterns. Like in Arabidopsis and other dicots, a subset of the rice HD-Zip I and II genes was found to be regulated by drought stress. We identified both drought-induced and drought-repressed HD-Zip genes and demonstrate that these genes are differentially regulated in drought-sensitive versus drought-tolerant rice cultivars. The drought-repressed HD-Zip family I gene, Oshox4, was selected for promoter-GUS analysis, showing that drought-responsiveness of Oshox4 is controlled by the promoter and that Oshox4 expression is predominantly vascular-specific. Loss-of-function analysis of Oshox4 revealed no specific phenotype, but overexpression analysis suggested a role for Oshox4 in elongation and maturation processes.

The Arabidopsis thaliana homeodomain leucine-zipper gene ATHB7 , which is active specifically under water deficit conditions, is proposed to act as a negative regulator of growth (Soderman et al ., 1996, Plant J. 10: 375 381; Hjellstrom et al ., 2003, Plant Cell Environ 26: 1127 1136). In this report we demonstrate that the paralogous gene, ATHB12 , has a similar expression pattern and function. ATHB12 ,like ATHB7 ,was up-regulated during water deficit conditions, the up-regulation being dependent on abscisic acid (ABA) and on the activity of the Ser/Thr phosphatases ABI1 and ABI2. Plants that are mutant for ATHB12 , as a result of T-DNA insertions in the ATHB12 gene, showed a reduced sensitivity to ABA in root elongation assays, whereas transgenic Arabidopsis plants expressing ATHB12 and/or ATHB7 as driven by the CaMV 35S promoter were hypersensitive in this response compared to wild-type. High-level expression of either gene also resulted in a delay in inflorescence stem elongation growth and caused plants to develop rosette leaves with a more rounded shape, shorter petioles, and increased branching of the inflorescence stem. Transgenic Arabidopsis plants expressing the reporter gene uidA under the control of the ATHB12 promoter showed marker gene activity in axillary shoot primordia, lateral root primordia, inflorescence stems and in flower organs. Treatment of plants with ABA or water deficit conditions caused the activity of ATHB12 to increase in the inflorescence stem, the flower organs and the leaves, and to expand into the vasculature of roots and the differentiation/elongation zone of root tips. Taken together, these results indicate that ATHB12 and ATHB7 act to mediate a growth response to water deficit by similar mechanisms.

Homeodomain-leucine zipper (HD-Zip) proteins are putative transcription factors encoded by a class of recently discovered homeobox genes as yet found only in plants. This paper reports on the characterization of one of these genes, ATHB-7, in Arabidopsis thaliana. ATHB-7 transcripts were present in all organs of the plant at low levels, but expression was induced several-fold by water deficit, osmotic stress as well as by exogenous treatment with abscisic acid (ABA), a response being detectable at 10(-8) M and reaching a maximum at 10(-6) M ABA. The ATHB-7 transcript was detected within 30 min after treatment with ABA and the transcript level was rapidly reduced after removal of the hormone. The induction of ATHB-7 was shown to be mediated strictly via ABA, since no induction of ATHB-7 was detectable in the ABA-deficient mutant aba-3 subjected to drought treatment. Induction levels in two ABA-insensitive mutants abi2 and abi3 were similar to the wild-type response. In the abi1 mutant, however, induction was impaired as 100-fold higher concentrations of ABA were required for a maximum induction as compared with wild-type. In this mutant the ATHB-7 response was reduced also after drought and osmotic stress treatments. These results indicate that ATHB-7 is transcriptionally regulated in an ABA-dependent manner and may act in a signal transduction pathway which mediates a drought response and also includes ABI1.

A set of MADS-box genes in flowering plants encode transcription factors that control both flower meristem formation and organ identity in the developing flower. In this report we present the first documentation of the presence of MADS-box genes in a non-flowering seed plant, and indeed from a plant bearing truly unisexual reproductive axes. A MADS-box-specific screening of a cDNA library from immature female strobili of the conifer Norway spruce, Picea abies (L.) Karst, resulted in cDNA clones that correspond to three different deficiens-agamous-like (dal) genes, dal1, dal2 and dal3. In addition to the MADS box, the spruce genes contain a second sequence element conserved among angiosperm genes, the K box, which is located downstream to the MADS box. A phylogenetic analysis of the nucleotide sequences confirms common ancestry of the gene superfamily. dal1 is related to agl2, agl4 and agl6 from Arabidopsis thaliana, all genes with unknown functions, and is expressed in vegetative as well as reproductive shoots on the adult spruce tree. dal2 is sister to angiosperm genes that control the identity of sexual organs, and is expressed only in the developing male and female strobili. dal3 is related to the vegetatively expressed tomato gene tm3 and is transcribed in both vegetative and reproductive shoots. These results strongly suggest that the functional and structural complexity within the MADS-box superfamily of reproduction-control genes is an ancestral property of seed plants and not a novelty in the angiosperm lineage.