Change of Fate and Staminodial Laminarity as Potential Agents of Floral Diversification in the Zingiberales.

The evolution of floral morphology in the monocot order Zingiberales shows a trend in which androecial whorl organs are progressively modified into variously conspicuous "petaloid" structures with differing degrees of fertility. Petaloidy of androecial members results from extensive laminarization of an otherwise radially symmetric structure. The genetic basis of the laminarization of androecial members has been addressed through recent candidate gene studies focused on understanding the spatiotemporal expression patterns of genes known to be necessary to floral organ formation. Here, we explore the correlation between gene duplication events and floral and inflorescence morphological diversification across the Zingiberales by inferring ancestral character states and gene copy number using the most widely accepted phylogenetic hypotheses. Our results suggest that the duplication and differential loss of GLOBOSA (GLO) copies is correlated with a change in the degree of the laminarization of androecial members. We also find an association with increased diversification in most families. We hypothesize that retention of paralogs in flower development genes could have led to a developmental shift affecting androecial organs with potential adaptive consequences, thus favoring diversification in some lineages but not others.

Evidence for the involvement of Globosa-like gene duplications and expression divergence in the evolution of floral morphology in the Zingiberales.

SUMMARY: *The MADS box transcription factor family has long been identified as an important contributor to the control of floral development. It is often hypothesized that the evolution of floral development across angiosperms and within specific lineages may occur as a result of duplication, functional diversification, and changes in regulation of MADS box genes. Here we examine the role of Globosa (GLO)-like genes, members of the B-class MADS box gene lineage, in the evolution of floral development within the monocot order Zingiberales. *We assessed changes in perianth and stamen whorl morphology in a phylogenetic framework. We identified GLO homologs (ZinGLO1-4) from 50 Zingiberales species and investigated the evolution of this gene lineage. Expression of two GLO homologs was assessed in Costus spicatus and Musa basjoo. *Based on the phylogenetic data and expression results, we propose several family-specific losses and gains of GLO homologs that appear to be associated with key morphological changes. The GLO-like gene lineage has diversified concomitant with the evolution of the dimorphic perianth and the staminodial labellum. *Duplications and expression divergence within the GLO-like gene lineage may have played a role in floral diversification in the Zingiberales.

[Pollen morphology in species of Canna (Cannaceae), and systematics implications]. / Morfología del polen en especies de Canna (Cannaceae) y su implicancia sistemática.

The morphology of pollen grains of eight taxa of Canna, C. ascendens, C. coccinea, C compacta, C. glauca, C. indica, C. paniculata, C. variegatifolia and C. fuchsina, an unpublished new species, were studied using light and scanning electronic microscopes. We used the Wodehouse technique on samples of 20 grains per specimen to measure the intine with a light microscope; and the density of spines (in 400 microm2 fields) with scanning electronic microscopy. Pollen grains are spherical, echinate, omniaperturate. The sporoderm presents a very thin exine covering a thicker intine. Corrugate micro-perforate, sub-reticulate, rugate, rugulate, striate to folded, micro-striate, micro-granulate, and smooth types of the external surface of the sporoderm were found. The spines consist of exine, partially to completely covered by tryphine. The two-layered intine is the thicker part of the wall. Echinate ornamentation is a generic character in Canna, but size, surface and color of pollen walls, and density and shape of spines, are diagnostic for species. Pollen morphology supports the view of C. indica and C. coccinea as different species. Canna fuchsina grows in wild, dense colonies, in humid riverside forests from Buenos Aires and Santa Fe Provinces, Argentina; its characters suggest relationships with a not well known group of taxa, some of them hybrids, such as C. x generalis. However, as these plants showed normal, well formed grains, close to those of C. coccinea, that germinate over the stigmatic surfaces in fresh flowers, we decided to include their pollen in this study.

Morfología del polen en especies de Canna (Cannaceae) y su implicancia sistemática

Pollen morphology in species of Canna (Cannaceae), and systematics implications. The morphology of pollen grains of eight taxa of Canna, C. ascendens, C. coccinea, C. compacta, C. glauca, C. indica, C. paniculata, C. variegatifolia and C. fuchsina, an unpublished new species, were studied using light and scanning electronic microscopes. We used the Wodehouse technique on samples of 20 grains per specimen to measure the intine with a light microscope; and the density of spines (in 400μm2 fields) with scanning electronic microscopy. Pollen grains are spherical, echinate, omniaperturate. The sporoderm presents a very thin exine covering a thicker intine. Corrugate micro-perforate, sub-reticulate, rugate, rugulate, striate to folded, micro-striate, micro-granulate, and smooth types of the external surface of the sporoderm were found. The spines consist of exine, partially to completely covered by tryphine. The two-layered intine is the thicker part of the wall. Echinate ornamentation is a generic character in Canna, but size, surface and color of pollen walls, and density and shape of spines, are diagnostic for species. Pollen morphology supports the view of C. indica and C. coccinea as different species. Canna fuchsina grows in wild, dense colonies, in humid riverside forests from Buenos Aires and Santa Fe Provinces, Argentina; its characters suggest relationships with a not well known group of taxa, some of them hybrids, such as C. x generalis. However, as these plants showed normal, well formed grains, close to those of C. coccinea, that germinate over the stigmatic surfaces in fresh flowers, we decided to include their pollen in this study. Rev. Biol. Trop. 58 (1): 63-79. Epub 2010 March 01.

Se estudió la morfología de los granos de polen de ocho táxones de Canna, C. ascendens, C. coccinea, C. compacta, C. glauca, C. indica, C. paniculata, C. variegatifolia y C. fuchsina, nueva especie aún no descrita, que fueron estudiadas usando microscopio de luz y microscopio electrónico de barrido. Nosotros utilizamos la técnica de Wodehouse en muestras de 20 granos por espécimen para medir la intina con el microscopio de luz; y la densidad de espinas (en campos de 400μm2) con el microscopio electrσnico de barrido. Los granos de polen son esfιricos, equinados y omniaberturados. El esporodermo presenta una exina muy delgada cubriendo una intina gruesa. La superficie del esporodermo puede ser corrugada, microperforada, sub-reticulada, rugada, rugulada, plegada-estriada, micro-estriada, microgranulada o lisa. Las espinas están formadas por exina, cubiertas total o parcialmente por trifina. La intina esta formada por dos capas, es la parte más gruesa de la pared. La ornamentación equinada es un rasgo genérico en Canna, pero el tamaño, la superficie y el color del polen, y la densidad y forma de las espinas, son rasgos diagnósticos de las especies. La morfología del polen apoya el tratamiento de C. coccinea y C. indica como especies diferentes. Canna fuchsina crece formando densas colonias silvestres en selvas ribereñas húmedas de las provincias de Buenos Aires y Santa Fe, Argentina; sus características sugieren relaciones no muy bien entendidas en el grupo de taxones, algunos son híbridos tales como C. x generalis. Sin embargo, estas plantas muestran granos normales, bien formados, cercanos a los de C. coccinea, que germinan sobre el estigma de flores frescas, nosotros decidimos incluir su polen en este estudio.

Unraveling the evolutionary radiation of the families of the Zingiberales using morphological and molecular evidence.

The Zingiberales are a tropical group of monocotyledons that includes bananas, gingers, and their relatives. The phylogenetic relationships among the eight families currently recognized are investigated here by using parsimony and maximum likelihood analyses of four character sets: morphological features (1), and sequence data of the (2) chloroplast rbcL gene, (3) chloroplast atpB gene, and (4) nuclear 18S rDNA gene. Outgroups for the analyses include the closely related Commelinaceae + Philydraceae + Haemodoraceae + Pontederiaceae + Hanguanaceae as well as seven more distantly related monocots and paleoherbs. Only slightly different estimates of evolutionary relationships result from the analysis of each character set. The morphological data yield a single fully resolved most-parsimonious tree. None of the molecular datasets alone completely resolves interfamilial relationships. The analyses of the combined molecular dataset provide more resolution than do those of individual genes, and the addition of the morphological data provides a well-supported estimate of phylogenetic relationships: (Musaceae ((Strelitziaceae, Lowiaceae) (Heliconiaceae ((Zingiberaceae, Costaceae) (Cannaceae, Marantaceae))))). Evidence from branch lengths in the parsimony analyses and from the fossil record suggests that the Zingiberales originated in the Early Cretaceous and underwent a rapid radiation in the mid-Cretaceous, by which time most extant family lineages had diverged.

Ecology. Food fight drives evolution.

On page 441 of this issue, evolutionary biologists showcase the purple-throated carib hummingbird as a rare example of food supply--in this case, flower shape--spurring the evolution of a sexual dimorphism, or a feature that differs between males and females. On St. Lucia, an island in the West Indies, female caribs sport bills a third longer and twice as curved as those of their male counterparts--one of the most extreme bill differences between the sexes in any hummingbird species. In the paper, the researchers link these "whoppingly dimorphic bills" to the specific flowers the male and female caribs frequent.

Evidence for ecological causation of sexual dimorphism in a hummingbird.

Unambiguous examples of ecological causes of animal sexual dimorphism are rare. Here we present evidence for ecological causation of sexual dimorphism in the bill morphology of a hummingbird, the purple-throated carib. This hummingbird is the sole pollinator of two Heliconia species whose flowers correspond to the bills of either males or females. Each sex feeds most quickly at the flower species approximating its bill dimensions, which supports the hypothesis that floral specialization has driven the evolution of bill dimorphism. Further evidence for ecological causation of sexual dimorphism was provided by a geographic replacement of one Heliconia species by the other and the subsequent development of a floral dimorphism, with one floral morph matching the bills of males and the other of females.

Malvidin 3-rutinoside as the pigment responsible for bract color in Curcuma alismatifolia.

Malvidin 3-rutinoside was the only anthocyanin identified from pink bracts of Curcuma alismatifolia cultivars. The concentration of malvidin 3-rutinoside in three cultivars increased as the intensity of the pink color in the bracts increased.

The functional morphology of the petioles of the banana, Musa textilis.

Bananas are among the largest herbs in the world and their lightweight petioles hold up huge leaves. This study examined how the petioles manage to achieve adequate rigidity to do this, while allowing extensive and reversible reconfiguration in high winds. Morphological and anatomical examination of the petioles and leaves of Musa textilis suggested how these two apparently incompatible abilities are achieved. The hollow U-shaped section of the petiole and the longitudinal strengthening elements in its outer skin give it adequate rigidity, while its ventral curvature help support the leaf without the need for thick lateral veins. These features, however, also allow the petiole to reconfigure by twisting away from the wind, while the leaf can fold away. In addition, two sets of internal structures, longitudinal partitions and transverse stellate parenchyma plates, help prevent dorsoventral flattening, allowing the petiole to flex further away from the wind without buckling. These ideas were tested and verified by a range of mechanical tests. Simple four-point-bending and torsion tests showed that the petioles are indeed far more compliant in torsion than in bending. Axial bending tests and crushing tests showed that petioles could be flexed twice as far and were four times as resistant to dorsoventral flattening when intact than when the internal tissue is removed. The banana petiole, therefore, seems to be an excellent example of natural integrated mechanical design.