In most plants, sucrose is exported from source leaves to carbon-importing sink tissues to sustain their growth and metabolism. Apoplastic phloem-loading species require sucrose transporters (SUTs) to transport sucrose into the phloem. In many dicot plants, genetic and biochemical evidence has established that SUT1-type proteins function in phloem loading. However, the role of SUT1 in phloem loading in monocot plants is not clear since the rice (Oryza sativa) and sugarcane (Saccharum hybrid) SUT1 orthologues do not appear to function in phloem loading of sucrose. A SUT1 gene was previously cloned from maize (Zea mays) and shown to have expression and biochemical activity consistent with a hypothesized role in phloem loading. To determine the biological function of SUT1 in maize, a sut1 mutant was isolated and characterized. sut1 mutant plants hyperaccumulate carbohydrates in mature leaves and display leaf chlorosis with premature senescence. In addition, sut1 mutants have greatly reduced stature, altered biomass partitioning, delayed flowering, and stunted tassel development. Cold-girdling wild-type leaves to block phloem transport phenocopied the sut1 mutants, supporting a role for maize SUT1 in sucrose export. Furthermore, application of 14C-sucrose to abraded sut1 mutant and wild-type leaves showed that sucrose export was greatly diminished in sut1 mutants compared with wild type. Collectively, these data demonstrate that SUT1 is crucial for efficient phloem loading of sucrose in maize leaves.

Auxin plays a fundamental role in organogenesis in plants. Multiple pathways for auxin biosynthesis have been proposed, but none of the predicted pathways are completely understood. Here, we report the positional cloning and characterization of the vanishing tassel2 (vt2) gene of maize (Zea mays). Phylogenetic analyses indicate that vt2 is a co-ortholog of TRYPTOPHAN AMINOTRANSFERASE OF ARABIDOPSIS1 (TAA1), which converts Trp to indole-3-pyruvic acid in one of four hypothesized Trp-dependent auxin biosynthesis pathways. Unlike single mutations in TAA1, which cause subtle morphological phenotypes in Arabidopsis thaliana, vt2 mutants have dramatic effects on vegetative and reproductive development. vt2 mutants share many similarities with sparse inflorescence1 (spi1) mutants in maize. spi1 is proposed to encode an enzyme in the tryptamine pathway for Trp-dependent auxin biosynthesis, although this biochemical activity has recently been questioned. Surprisingly, spi1 vt2 double mutants had only a slightly more severe phenotype than vt2 single mutants. Furthermore, both spi1 and vt2 single mutants exhibited a reduction in free auxin levels, but the spi1 vt2 double mutants did not have a further reduction compared with vt2 single mutants. Therefore, both spi1 and vt2 function in auxin biosynthesis in maize, possibly in the same pathway rather than independently as previously proposed.

Patterning of the floral organs is exquisitely controlled and executed by four classes of homeotic regulators. Among these, the class B and class C floral homeotic regulators are of central importance as they specify the male and female reproductive organs. Inappropriate induction of the class B gene APETALA3 (AP3) and the class C gene AGAMOUS (AG) causes reduced reproductive fitness and is prevented by polycomb repression. At the onset of flower patterning, polycomb repression needs to be overcome to allow induction of AP3 and AG and formation of the reproductive organs. We show that the SWI2/SNF2 chromatin-remodeling ATPases SPLAYED (SYD) and BRAHMA (BRM) are redundantly required for flower patterning and for the activation of AP3 and AG. The SWI2/SNF2 ATPases are recruited to the regulatory regions of AP3 and AG during flower development and physically interact with two direct transcriptional activators of class B and class C gene expression, LEAFY (LFY) and SEPALLATA3 (SEP3). SYD and LFY association with the AP3 and AG regulatory loci peaks at the same time during flower patterning, and SYD binding to these loci is compromised in lfy and lfy sep3 mutants. This suggests a mechanism for SWI2/SNF2 ATPase recruitment to these loci at the right stage and in the correct cells. SYD and BRM act as trithorax proteins, and the requirement for SYD and BRM in flower patterning can be overcome by partial loss of polycomb activity in curly leaf (clf) mutants, implicating the SWI2/SNF2 chromatin remodelers in reversal of polycomb repression. P lant development occurs largely postembryonically (1), and, as a consequence, many cell-fate choices do not take place until long after embryogenesis. One example is flower development; in the rapid-flowering winter annual Arabidopsis the first flowers are formed 1 mo to 1 y after germination (2). Precocious activation of the floral homeotic genes required for flower patterning results in pleiotropic defects including poor seed set and is prevented by chromatin repression, which is faithfully inherited throughout cell divisions until the first flowers are formed (3-7). The repressive chromatin needs to be erased for flower patterning to be initiated in flower primordia. For class B genes, such as APETALA3 (AP3), which are required for correct patterning of the showy petals and the male reproductive organs, activation of gene expression occurs in late stage 2 flower primordia in the cells that will give rise to whorls 2 and 3 of the flower (6). For class C gene AGAMOUS (AG), which is required for patterning both the male and the female reproductive organs, induction is observed in early stage 3 flowers in the cells that will give rise to whorls 3 and 4 (6).The mitotically heritable chromatin repression of AP3 and AG before flower formation is achieved by two polycomb complexes: polycomb repressive complex 1 (PRC1) and PRC2. PRC2 is responsible for trimethylation of lysine 27 of histone H3 (H3K27me3) (7,8). Two putative H3K27 methyltransferases and PRC2 complex components, SWINGER and ...

A dramatic change in agricultural crops is needed in order to keep pace with the demands of an increasing human population, exponential need for renewable fuels, and uncertain climatic changes. Grasses make up the vast majority of agricultural commodities. How these grasses capture, transport, and store carbohydrates underpins all aspects of crop productivity. Sink-source dynamics within the plant direct how much, where, and when carbohydrates are allocated, as well as determine the harvestable tissue. Carbohydrate partitioning can limit the yield capacity of these plants, thus offering a potential target for crop improvement. Grasses have the ability to buffer this sink-source interaction by transiently storing carbohydrates in stem tissue when production from the source is greater than whole-plant demand. These reserves improve yield stability in grain crops by providing an alternative source when photosynthetic capacity is reduced during the later phases of grain filling, or during periods of environmental and biotic stresses. Domesticated grasses such as sugarcane and sweet sorghum have undergone selection for high accumulation of stem carbohydrates, which serve as the primary sources of sugars for human and animal consumption, as well as ethanol production for fuel. With the enormous expectations placed on agricultural production in the near future, research into carbohydrate partitioning in grasses is essential for maintaining and increasing yields in grass crops. This review highlights the current knowledge of non-structural carbohydrate dynamics in grass stems and discusses the impacts of stem reserves in essential agronomic grasses.

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