Plant development. Genetic control of distal stem cell fate within root and embryonic meristems.

The root meristem consists of populations of distal and proximal stem cells and an organizing center known as the quiescent center. During embryogenesis, initiation of the root meristem occurs when an asymmetric cell division of the hypophysis forms the distal stem cells and quiescent center. We have identified NO TRANSMITTING TRACT (NTT) and two closely related paralogs as being required for the initiation of the root meristem. All three genes are expressed in the hypophysis, and their expression is dependent on the auxin-signaling pathway. Expression of these genes is necessary for distal stem cell fate within the root meristem, whereas misexpression is sufficient to transform other stem cell populations to a distal stem cell fate in both the embryo and mature roots.

HALF FILLED promotes reproductive tract development and fertilization efficiency in Arabidopsis thaliana.

Successful fertilization in angiosperms requires the growth of pollen tubes through the female reproductive tract as they seek out unfertilized ovules. In Arabidopsis, the reproductive tract begins with the stigma, where pollen grains initially adhere, and extends through the transmitting tract of the style and ovary. In wild-type plants, cells within the transmitting tract produce a rich extracellular matrix and undergo programmed cell death to facilitate pollen movement. Here, we show that the HAF, BEE1 and BEE3 genes encode closely related bHLH transcription factors that act redundantly to specify reproductive tract tissues. These three genes are expressed in distinct but overlapping patterns within the reproductive tract, and in haf bee1 bee3 triple mutants extracellular matrix formation and cell death fail to occur within the transmitting tract. We used a minimal pollination assay to show that HAF is necessary and sufficient to promote fertilization efficiency. Our studies further show that HAF expression depends on the NTT gene and on an auxin signaling pathway mediated by the ARF6, ARF8 and HEC genes.

The formation and function of the female reproductive tract in flowering plants.

In angiosperms, sexual reproduction requires a sperm cell, contained within a pollen tube, to fertilize the egg cell. The pollen tubes are capable of growth but have a difficult journey, as egg cells are buried within the ovary of the carpel. Several tissues, known collectively as the reproductive tract, develop within the carpel to facilitate the journey of the pollen tube. The genes involved in the formation and function of the reproductive tract have largely remained a mystery but are crucial for successful fertilization. This review summarizes recent advances in our understanding of the genetic control of reproductive tract development.

The NTT gene is required for transmitting-tract development in carpels of Arabidopsis thaliana.

BACKGROUND: The majority of pollen-tube growth in Arabidopsis occurs in specialized tissue called the transmitting tract. Little is currently known about how the transmitting tract functions because of a lack of mutants affecting its development. We have identified such a mutant and we used it to investigate aspects of pollen-tube growth. RESULTS: Reverse genetics was used to identify mutations in a gene, No Transmitting Tract (NTT), encoding a C2H2/C2HC zinc finger transcription factor specifically expressed in the transmitting tract. The ntt mutants have a negative effect on transmitting-tract development. Stage-specific analysis of transmitting-tract development was carried out and was correlated with investigations of pollen-tube behavior. In ntt mutants, pollen tubes grow more slowly and/or terminate prematurely, and lateral divergence is accentuated over apical-to-basal movement. Normal transmitting-tract development is shown to involve a process of programmed cell death (PCD) that is facilitated by, but does not depend upon, pollination. CONCLUSIONS: This is the first report of a gene that is specifically required for transmitting-tract development in Arabidopsis. Mutations in NTT cause reduced fertility by severely inhibiting pollen-tube movement. The data support the idea that the function of the transmitting tract is to increase fertilization efficiency, particularly in the lower half of the ovary. This occurs by facilitating pollen-tube growth through differentiation and then death of transmitting-tract cells.

CINCINNATA controls both cell differentiation and growth in petal lobes and leaves of Antirrhinum.

To understand how differentiation and growth may be coordinated during development, we have studied the action of the CINCINNATA (CIN) gene of Antirrhinum. We show that in addition to affecting leaf lamina growth, CIN affects epidermal cell differentiation and growth of petal lobes. Strong alleles of cin give smaller petal lobes with flat instead of conical cells, correlating with lobe-specific expression of CIN in the wild type. Moreover, conical cells at the leaf margins are replaced by flatter cells, indicating that CIN has a role in cell differentiation of leaves as well as petals. A weak semidominant cin allele affects cell types mainly in the petal but does not affect leaf development, indicating these two effects can be separated. Expression of CIN correlates with expression of cell division markers, suggesting that CIN may influence petal growth, directly or indirectly, through effects on cell proliferation. For both leaves and petals, CIN affects growth and differentiation of the more distal and broadly extended domains (leaf lamina and petal lobe). However, while CIN promotes growth in petals, it promotes growth arrest in leaves, possibly because of different patterns of growth control in these systems.

Genetic control of surface curvature.

Although curvature of biological surfaces has been considered from mathematical and biophysical perspectives, its molecular and developmental basis is unclear. We have studied the cin mutant of Antirrhinum, which has crinkly rather than flat leaves. Leaves of cin display excess growth in marginal regions, resulting in a gradual introduction of negative curvature during development. This reflects a change in the shape and the progression of a cell-cycle arrest front moving from the leaf tip toward the base. CIN encodes a TCP protein and is expressed downstream of the arrest front. We propose that CIN promotes zero curvature (flatness) by making cells more sensitive to an arrest signal, particularly in marginal regions.