Alterations of CorTFL1 and CorAP1 expression correlate with major evolutionary shifts of inflorescence architecture in Cornus (Cornaceae) - a proposed model for variation of closed inflorescence forms.

TFL1-, AP1- and LFY-like genes are known to be key regulators of inflorescence development. However, it remains to be tested whether the evolutionary modifications of inflorescence morphology result from shifts in their expression patterns. We compared the spatiotemporal expression patterns of CorTFL1, CorAP1 and CorLFY in six closely related Cornus species that display four types of closed inflorescence morphology using quantitative real-time polymerase chain reaction (qRT-PCR) and RNA in situ hybridization. Character mapping on the phylogeny was conducted to identify evolutionary changes and to assess the correlation between changes in gene expression and inflorescence morphology. Results demonstrated variation of gene expression patterns among species and a strong correlation between CorTFL1 expression and the branch index of the inflorescence type. Evolutionary changes in CorTFL1 and CorAP1 expression co-occurred on the phylogeny with the morphological changes underpinning inflorescence divergence. The study found a clear correlation between the expression patterns of CorTFL1 and CorAP1 and the inflorescence architecture in a natural system displaying closed inflorescences. The results suggest a role for the alteration in CorTFL1 and CorAP1 expression during the evolutionary modification of inflorescences in Cornus. We propose that a TFL1-like and AP1-like gene-based model may explain variation of closed inflorescences in Cornus and other lineages.

Evolution of bract development and B-class MADS box gene expression in petaloid bracts of Cornus s. l. (Cornaceae).

Despite increasing interest in the molecular mechanisms of floral diversity, few studies have investigated the developmental and genetic bases of petaloid bracts. This study examined morphological patterns of bract initiation and expression patterns of B-class MADS-box genes in bracts of several Cornus species. We suggest that petaloid bracts in this genus may not share a single evolutionary origin. Developmental pathways of bracts and spatiotemporal expression of B-class genes in bracts and flowers were examined for four closely related dogwood species. Divergent morphological progressions and gene expression patterns were found in the two sister lineages with petaloid bracts, represented by Cornus florida and Cornus canadensis. Phylogeny-based analysis identified developmental and gene expression changes that are correlated with the evolution of petaloid bracts in C. florida and C. canadensis. Our data support the existence of independent evolutionary origins of petaloid bracts in C. canadensis and C. florida. Additionally, we suggest that functional transference within B-class gene families may have contributed to the origin of bract petaloidy in C. florida. However, the underlying mechanisms of petaloid bract development likely differ between C. florida and C. canadensis. In the future this hypothesis can be tested by functional analyses of Cornus B-class genes.

Plasticity in leaf-area density within the crown of Aucuba japonica growing under different light levels.

We assessed leaf-area density (LAD; m(2) m(-3)) within the crown of Aucuba japonica (Cornaceae) growing under different light regimes and analyzed the components of crown architecture that most influenced variation in LAD. At a whole-crown level, extension-unit (EU) density (EUs/m(3)) had the greatest impact on LAD. The number of leaves per unit EU length and EU length had a wide range of impacts depending on the degree of crowding of foliage on the EU. Leaf size had a lesser impact on LAD. LAD was higher in the uppermost crown and declined towards the base. The non-uniformity of LAD among crown layers was much greater under high irradiance. Individuals under high irradiance achieved greater LAD by increased branching, well-marked EU dimorphism and a larger number of leaves per unit EU length; the reverse was true for the individuals under low irradiance. We identified two distinct modes of growth response to light regime. Under high irradiance, individuals responded by differential growth between the layers of crowns with the lower crown suppressed and growth in the upper crown increased. Conversely, shaded individuals did not respond by differential growth between crown layers.

Shoot morphology of Aucuba japonica incurred by anisophylly: ecological implications.

Anisophylly, having leaves different in size and/or shape, was quantified in adult Aucuba japonica and simulations were carried out to evaluate the effects of anisophylly on the extent of self-shading at the single-shoot level as well as at the whole-canopy level. Clear anisophylly was observed in the individual after switching from the single-stemmed juvenile stage to the multi-stemmed adult stage. In such plants, leaf area in the canopy abruptly increased. The effective display of adult foliage involved a variety of morphological changes in addition to anisophylly, most prominently reduction in leaf size compared to juveniles. The simulation results indicate that diversity of leaf size and shape is an effective means of minimizing self-shading as well as allowing the efficient exploitation of a larger canopy volume in adult plants. Anisophylly also increased the biomass use efficiency of individual plants at maturity. Taken together, having diverse leaf forms is superior to having a single leaf form for maximizing area acquisition and for efficiently filling the acquired area. We therefore conclude that the anisophylly expressed in A. japonica is adaptive.

Flower bud abortion influences clonal growth and sexual dimorphism in the understorey dioecious shrub Aucuba japonica (Cornaceae).

Sexual differences were investigated to determine the significance of flower bud abortion in the dioecious shrub Aucuba japonica Thunb. The mean number of flowers per inflorescence and the mean number of flowering inflorescences (as opposed to aborted inflorescences) per individual were greater in males than in females in 1997 and 1998. Reproductive investment by males was 0.4-times (1997) and 1.4-times (1998) that by females. In addition, females aborted 30.9% (1997) and 42.7% (1998) of their total flower buds without blooming, whereas no male flower buds aborted. One of the architectural traits of this shrub is that in the year that a flower bud is produced at the shoot apex, the shoot will branch into two or more shoots. Thus, there was less sexual difference in the number of current shoots per individual than there was in the number of flowering inflorescences. The relationship between annual growth and reproduction, and the probability of reproduction in the following year, suggested that the higher investment in female reproduction was manifested as a cost for reproductive frequency rather than as a cost for annual growth. The spatial distribution of both males and females was clumped, which may be the result of clonal growth. In addition, overall sex ratios were not skewed and the number of sprouts did not differ significantly between sexes. These results suggested that flower bud abortion by females might reduce sexual dimorphism in terms of clonal growth.