Insights into the functional and genetic basis of heteranthery in Arthrostemma ciliatum Pav. ex D.Don (Melastomataceae)

BACKGROUND: Melastomes are well known for their striking diversity in stamen morphologies mostly adapted to buzz pollination by bees. The variously modified connective appendages and heteranthery in the family have fascinated botanists for more than two centuries and a variety of functions associated with pollination have been discovered for these staminal traits over the years. The repeated evolutionary shifts in staminal traits have been linked to pollinator shifts, likely contributing to diversification in the family. The evolutionary lability of staminal traits, especially the connective morphology, led us to hypothesize that they might be controlled by relatively simple genetic mechanisms and we here take the first steps to test this hypothesis by using a comparative transcriptomics approach with Arthrostemma ciliatum as our model. We also tested the functional significance of heteranthery and whether the classical division of labour hypothesis holds true for this species by comparing the number, size and viability of pollen in the two stamen types. RESULTS: Staminal development of this species was studied and suitable stages for transcriptome comparisons were identified. Differential expression analyses between the morphologically distinct stamen whorls at four developmental stages showed the differential expression of several transcripts involved in stamen development/elongation. Pollen comparisons between the two whorls showed that the antepetalous/inner whorl stamens have a significantly higher number of pollen and higher germination rates while the antesepalous/outer whorl stamens have significantly larger pollen. CONCLUSIONS: We identified Jasmonate and Gibberellin signalling pathway genes (JAZ, GID1, DELLA and ARF homologs), EPF/EPFL family genes, autophagy related genes (VPE homologs) and S Locus ELF homologs as putative candidates, likely involved in causing staminal dimorphism in A. ciliatum. Our results indicate that, for the heterantherous morph of this species, the shorter stamens (antepetalous/inner whorl) have both pollinating and feeding functions contradicting the division of labour theory. We also report the possible existence of heterostyly in A. ciliatum as an outcrossing mechanism.