Metamerism,heterochrony, and inflorescence morphology of the Pithecellobium-complex (Leguminosae: Mimosoideae: Ingeae)

The diverse inflorescence morphology of species in the Pithecellobium-complex is shown to be a result of: 1) the organization of the components of the inflorescence and their relative positions; 2) the hierarchical relationship of the axes of the inflorescences and the position they assume in total tree architecture; and 3) heterochronic development of the components of the inflorescence. It is shown that the typological system of nomenclature of inflorescences leads to false assumptions of homology and therefore must be discarded. The morphology of inflorescences is discussed in terms of metamerism, and the term Repeating Growth Unit (RGU) is introduced and is defined as the smallest complete sequence of metamers produced by a meristem. A module is defined as the sequence of RGUs produced by a meristem. An inflorescence is defined as that sequence of metamers in an RGU which participates in the production and/or presentation of flowers and fruit. Heterochrony, proleptic and sylleptic buds, and shoot dimorphism are discussed and their role in modifying inflorescence morphology in the Pithecellobium-complex is illustrated. Examples from the Pithecellobium-complex are provided which demonstrate the various modifications of the inflorescence that result from the interaction of these various phenomena.     

Molecular phylogenetics of Acacia (Fabaceae: Mimosoideae) based on the chloroplast MATK coding sequence and flanking TRNK intron spacer regions

The tribe Acacieae (Fabaceae: Mimosoideae) contains two genera, the monotypic African Faidherbia and the pantropical Acacia, which comprise about 1200 species with over 950 confined to Australia. As currently recognized, the genus Acacia is subdivided into three subgenera: subg. Acacia, subg. Aculeiferum, and the predominantly Australian subg. Phyllodineae. Morphological studies have suggested the tribe Acacieae and genus Acacia are artificial and have a close affinity to the tribe Ingeae. Based on available data there is no consensus on whether Acacia should be subdivided. Sequence analysis of the chloroplast trnK intron, including the matK coding region and flanking noncoding regions, indicate that neither the tribe Acacieae nor the genus Acacia are monophyletic. Two subgenera are monophyletic; section Filicinae of subgenus Aculeiferum does not group with taxa of the subgenus. Section Filicinae, eight Ingeae genera, and Faidherbia form a weakly supported paraphyletic grade with respect to subg. Phyllodineae. Acacia subg. Aculeiferum (s. s.) is sister to the grade. These data suggest that characters currently used to differentiate taxa at the tribal, generic, and subgeneric levels are polymorphic and homoplasious in cladistic analyses.     

A plastid DNA phylogeny of the genus Acacia Miller (Acacieae, Leguminoseae)

Past classifications of the tribe Acacieae Rchb. are outlined and the confusion concerning the relationships of the three subgenera of Acacia Mill. are highlighted. A plastid DNA analysis of Acacieae shows that the genus Acacia is not monophyletic. Furthermore subgenera Acacia Vassal and Aculeiferum Vassal are sister taxa and neither appear closely related to subgenus Phyllodineae (DC.) Ser. Subgenera Acacia and Aculeiferum form a clade that is basal to a well-supported clade consisting of tribe Ingeae Benth. taxa, Faidherbia albida (Del.) A. Chev. and subgenus Phyllodineae. The series of relationships suggested by the cpDNA data contradicts previous investigations of the tribe. Possible explanations of this conflict are explored, and the taxonomic implications of the plastid DNA data set are considered.     

POLLEN ULTRASTRUCTURE OF THE TRIBE INGEAE (MIMOSOIDEAE: LEGUMINOSAE)

All genera within the Ingeae, excluding Wallaceodendron, were examined with the transmission electron microscope. Thin sections reveal two pollen types (Types I and II) distinguished primarily by differences in polyad cohesion and ektexine organization. Type I polyads (only eight-grained species of Calliandra) are calymmate and the ektexine of individual cells is continuous around the grain, organized into a thin, foraminate tectum, irregularly shaped, often basally flared, foraminate columellae and thin, discontinuous foot layer. Type II polyads (16-grained species of Calliandra and remaining Ingeae) are predominantly acalymmate with individual grains typically free from one another or rarely, partially calymmate, i.e., individual grains show limited forms of attachment through small endexinous bridges (Pithecellobium latifolium [Zygia], Lysiloma) or localized appression of adjacent endexines (Pithecellobium daulense [Cathormion]). The adhesion of individual grains through localized fusion of lateral-distal and proximal ektexine in Enterolobium is unique among the partially calymmate Type II polyads. Ektexine in Type II polyads, largely restricted to the distal face, is composed of a thick, channeled tectum, granular interstitium and when present, thin discontinuous foot layer. Lateral-distal and proximal areas exhibit only endexine and, occasionally, a foot layer. The occurrence of nondistal ektexine is restricted to Enterolobium. The pollen data suggest that the acalymmate Ingeae polyads composed of grains with porate apertures, thick, highly channeled tectum, granular interstitium and lack of, or greatly reduced foot layer, are clearly derived within the Mimosoideae. Type I calymmate polyads appear to be independently derived. Ultrastructural data suggest that the Ingeae, excluding the eight-grained Calliandra species, represent a natural grouping with a close affinity to the Acacieae.     

HETEROMORPHIC FLOWER DEVELOPMENT IN NEPTUNIA PUBESCENS,A MIMOSOID LEGUME

The characteristic of heteromorphic inflorescences in some mimosoid legumes such as Neptunia is a puzzling one which can be approached developmentally. Each spicate inflorescence of Neptunia pubescens includes three types of flowers: perfect in the upper half, functionally male just below the middle, and sterile or neuter at the base. Developmental studies of the inflorescence show that order of initiation of bracts on the inflorescence is acropetal, but that order of subsequent development of flowers is both acropetal and basipetal on the axis. Bract growth and initiation of the axillary floral apices at the base are inhibited or retarded, while those in the middle and upper levels continue development without interruption. The three types of floral primordia are similar during initiatory stages of organ formation and through early development. At mid-development, differences arise in floral symmetry, petal form, stamen form, and size and shape of the carpel. The functionally male flowers become strongly dorsiventral and zygomorphic while the other two morphs remain actinomorphic or nearly so. Heteromorphy arises from a combination of early suppression of organogeny plus mid-stage innovations of zygomorphy and lateral expansion of stamen primordia. These divergent developmental pathways in one inflorescence can be interpreted in part using Gould's concept of heterochrony: changes in timing of developmental events to produce different structures. Other changes in Neptunia cannot be explained by this concept, however; such changes as omission of processes (i.e., meiosis) in some organs, or addition of processes not normally present (i.e., blade formation in stamen primordia which become staminodia). It is becoming evident from work on this and other legume flowers that actual loss of organs is rare, compared to initiation followed by suppression or modification.     

<Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> as a model for gelatinous fiber formation

Trees and herbaceous plants continuously monitor their position to maintain vertical stem growth and regulate branch orientation. When orientation is altered from the vertical, they form a special type of wood called reaction wood that differs chemically and structurally from normal wood and forces reorientation of the organ or whole plant. The reaction wood of dicotyledons is called tension wood and is characterized by nonlignified gelatinous fibers. The altered chemical and mechanical properties of tension wood reduce wood quality and represent a major problem for the timber and pulping industries. Repeated clipping of the emerging inflorescence stems of Arabidopsis thaliana augments wood formation in organs, including those inflorescence stems that are allowed to develop later. Gravistimulation of such inflorescence stems induces tension wood formation, allowing the use of A. thaliana for a molecular and genetic analysis of the mechanisms of tension wood formation.     

Flowering and determinacy in maize

All plant organs are produced by meristems, groups of stem cells located in the tips of roots and shoots. Indeterminate meristems make an indefinite number of organs, whereas determinate meristems are consumed after making a specific number of organs. Maize is an ideal system to study the genetic control of meristem fate because of the contribution from determinate and indeterminate meristems to the overall inflorescence. Here, the latest work on meristem maintenance and organ specification in maize is reviewed. Genetic networks, such as the CLAVATA components of meristem maintenance and the ABC programme of flower development, are conserved between grasses and eudicots. Maize and rice appear to have conserved mechanisms of meristem maintenance and organ identity. Other pathways, such as sex determination, are likely to be found only in maize with its separate male and female flowers. A rich genetic history has resulted in a large collection of maize mutants. The advent of genomic tools and synteny across the grasses now permits the isolation of the genes behind inflorescence architecture and the ability to compare function across the Angiosperms.     

Genetic characterization and floral development of female sterile and stubby head, two aposporous mutants of pearl millet

 Apomixis has never been reported in natural populations of pearl millet [Pennisetum glaucum (L.) R.Br.], although many wild relatives of pearl millet are obligate or facultative aposporous apomicts. Four-nucleate aposporous embryo sacs are formed from somatic cells of the nucellus that do not undergo meiosis. Two mutants of pearl millet, female sterile (fs) and stubby head, have two developmental characteristics in common: a significant reduction in head length compared with the wild-type and the formation of aposporous embryo sacs. Reproductive development in fs and stubby head mutants was examined in depth because of the potential for illuminating basic cellular or developmental factors that may function to alter embryo sac development. Genetic analysis of stubby head showed that this phenotype is conferred by genes at two loci linked in coupling within 29 cM. Crosses between fs and stubby head mutants showed that, despite the similarities in phenotypes, the mutations are at different loci. The mutants differ from wild-type in their inflorescence structure from the time of initiation of spikelet primordia through terminal differentiation of the ovule. Both mutations could be categorized as meristic, since a change in inflorescence branch or organ number was common and gynoecium development varied. We speculate that heterochronic development of the floral meristem and organ initiation/specification programs may be the underlying mechanism for phenotypic changes in these mutants throughout the floral phase. Received: 25 October 1996 / Accepted: 13 March 1997     

Racemose inflorescences of monocots: structural and morphogenetic interaction at the flower/inflorescence level

Background

Understanding and modelling early events of floral meristem patterning and floral development requires consideration of positional information regarding the organs surrounding the floral meristem, such as the flower-subtending bracts (FSBs) and floral prophylls (bracteoles). In common with models of regulation of floral patterning, the simplest models of phyllotaxy consider only unbranched uniaxial systems. Racemose inflorescences and thyrses offer a useful model system for investigating morphogenetic interactions between organs belonging to different axes.

Scope

This review considers (1) racemose inflorescences of early-divergent and lilioid monocots and their possible relationship with other inflorescence types, (2) hypotheses on the morphogenetic significance of phyllomes surrounding developing flowers, (3) patterns of FSB reduction and (4) vascular patterns in the primary inflorescence axis and lateral pedicels.

Conclusions

Racemose (partial) inflorescences represent the plesiomorphic condition in monocots. The presence or absence of a terminal flower or flower-like structure is labile among early-divergent monocots. In some Alismatales, a few-flowered racemose inflorescence can be entirely transformed into a terminal ‘flower’. The presence or absence and position of additional phyllomes on the lateral pedicels represent important taxonomic markers and key features in regulation of flower patterning. Racemose inflorescences with a single floral prophyll are closely related to thyrses. Floral patterning is either unidirectional or simultaneous in species that lack a floral prophyll or possess a single adaxial floral prophyll and usually spiral in the outer perianth whorl in species with a transversely oriented floral prophyll. Inhibitory fields of surrounding phyllomes are relevant but insufficient to explain these patterns; other important factors are meristem space economy and/or the inhibitory activity of the primary inflorescence axis. Two patterns of FSB reduction exist in basal monocots: (1) complete FSB suppression (cryptic flower-subtending bract) and (2) formation of a ‘hybrid’ organ by overlap of the developmental programmes of the FSB and the first abaxial organ formed on the floral pedicel. FSB reduction affects patterns of interaction between the conductive systems of the flower and the primary inflorescence axis.     

Comparative organ differentiation during early life stages of marine fish

The basic developmental mechanisms of teleosts are similar, but there are differences with respect to the timing of developmental events. These events are controlled by genetic and environmental factors. Direct comparisons of organogenesis are complicated due to large variations in egg sizes and incubation temperatures between species. But in general, cultivated small marine pelagic fish larvae originating from rather small eggs (like gadoids, flatfishes, sparids) hatch with a relatively large yolk sac, a larval finfold and subdermal space and under-developed organs. Developmental status at hatch differs between species and the duration of the yolk sac period varies. Main organs and organ systems become functional by first feeding and differentiate during the larval stage and metamorphosis. Species developing directly via large yolk-rich eggs and a long incubation period have a juvenile like morphology and organ functionality at first feeding, sometimes immediately after hatch (like wolffishes). Histomorphological and cell- or organ functional studies of developing embryos and larvae of cultivated species constitute basic information for understanding species-specific events, of utmost importance for improving production protocols. Information is still lacking on early functionality of endocrine and immunocompetent tissues and organs, areas that deserve future focus.     

Direct and indirect selection on floral pigmentation by pollinators and seed predators in a color polymorphic South African shrub

The coexistence of different color morphs is often attributed to variable selection pressures across space, time, morph frequencies, or selection agents, but the routes by which each morph is favored are rarely identified. In this study we investigated factors that influence floral color polymorphisms on a local scale in Protea, within which approximately 40% of species are polymorphic. Previous work shows that seed predators and reproductive differences likely contribute to maintaining polymorphism in four Protea species. We explored whether selection acts directly or indirectly on floral color in two populations of Protea aurea, using path analysis of pollinator behavior, nectar production, seed predation, color, morphology, and maternal fecundity fitness components. We found that avian pollinators spent more time on white morphs, likely due to nectar differences, but that this had no apparent consequences for fecundity. Instead, the number of flowers per inflorescence underpinned many of the reproductively important differences between color morphs. White morphs had more flowers per inflorescence, which itself was positively correlated with nectar production, seed predator occurrence, and total long-term seed production. The number of seeds per plant to survive predation, in contrast, was not directly associated with color or any other floral trait. Thus, although color differences may be associated with conflicting selection pressures, the selection appears to be associated with the number of flowers per inflorescence and its unmeasured correlates, rather than with inflorescence color itself.     

Cotyledon architecture and anatomy in the Acacieae (Leguminosae: Mimosoideae)

Cotyledon size, shape, venation pattern and anatomy have been investigated in Faidherbia albida and 152 species of Acacia representing the three subgenera Acacia, Aculeiferum and Heterophyllum. Cell volumes of epidermis, palisade and storage tissue, stomatal frequency, stomatal index and frequency of stomatal types have been determined for F. albida and 12 species from each subgenus. The data obtained support the recognition of the subgenera of Acacia as separate taxa but provide no indication of their status. The evidence from cotyledons also supports the separation of Faidherbia from Acacia , and the amalgamation of the Acacieae and Ingeae.     

A putative lipase gene EXTRA GLUME1 regulates both empty-glume fate and spikelet development in rice

Recent studies have shown that molecular control of inner floral organ identity appears to be largely conserved between monocots and dicots, but little is known regarding the molecular mechanism underlying development of the monocot outer floral organ, a unique floral structure in grasses. In this study, we report the cloning of the rice EXTRA GLUME1 ( EG1 ) gene, a putative lipase gene that specifies empty-glume fate and floral meristem determinacy. In addition to affecting the identity and number of empty glumes, mutations in EG1 caused ectopic floral organs to be formed at each organ whorl or in extra ectopic whorls. Iterative glume-like structures or new floral organ primordia were formed in the presumptive region of the carpel, resulting in an indeterminate floral meristem. EG1 is expressed strongly in inflorescence primordia and weakly in developing floral primordia. We also found that the floral meristem and organ identity gene OsLHS1 showed altered expression with respect to both pattern and levels in the eg1 mutant, and is probably responsible for the pleiotropic floral defects in eg1 . As a putative class III lipase that functionally differs from any known plant lipase, EG1 reveals a novel pathway that regulates rice empty-glume fate and spikelet development.     

The tympanal hearing organ of the parasitoid fly Ormia ochracea (Diptera, Tachinidae, Ormiini)

Tympanate hearing has evolved in at least 6 different orders of insects, but had not been reported until recently in the Diptera. This study presents a newly discovered tympanal hearing organ, in the parasitoid tachinid fly, Ormia ochracea. The hearing organ is described in terms of external and internal morphology, cellular organization of the sensory organ and preliminary neuroanatomy of the primary auditory afferents. The ear is located on the frontal face of the prothorax, directly behind the head capsule. Conspicuously visible are a pair of thin cuticular membranes specialized for audition, the prosternal tympanal membranes. Directly attached to these membranes, within the enlarged prosternal chamber, are a pair of auditory sensory organs, the bulbae acusticae. These sensory organs are unique among all auditory organs known so far because both are contained within an unpartitioned acoustic chamber. The prosternal chamber is connected to the outside by a pair of tracheae. The cellular anatomy of the fly's scolopophorous organ was investigated by light and electron microscopy. The bulba acustica is a typical chordotonal organ and it contains approximately 70 receptor cells. It is similar to other insect sensory organs associated with tympanal ears. The similarity of the cellular organization and tympanal morphology of the ormiine ear to the ears of other tympanate insects suggests that there are potent constraints in the design features of tympanal hearing organs, which must function to detect high frequency auditory signals over long distances. Each sensory organ is innervated by a branch of the frontal nerve of the fused thoracic ganglia. The primary auditory afferents project to each of the pro-, meso-, and metathoracic neuropils. The fly's hearing organ is sexually dimorphic, whereby the tympanal membranes are larger in females and the spiracles larger in males. The dimorphism presumably reflects differences in the acoustic behavior in the two sexes.     

The genesis of neural crest and epidermal placodes: a reinterpretation of vertebrate origins

Vertebrate body organization differs from that of other chordates in a large number of derived features that involve all organ systems. Most of these features arise embryonically from epidermal placodes, neural crest, and a muscularized hypomere. The developmental modifications were associated with a shift from filter-feeding to more active predation, which established advantages for improved gas exchange and distribution. Active predation involved more efficient patterns of locomotion and led to a major reorganization of the pharynx, to elaboration of the circulatory, digestive, and nervous systems, and to special sense organs. Most of the organs that derive from epidermal placodes and neural crest may have arisen phylogentically from epidermal nerve plexus of earlier chordates. Supportive tissues such as cartilage, bone, dentine, and enamel-like tissues probably arose in association with several of the new vertebrate sense organs and only secondarily provided mechanical support. The development of armor appears to have occurred late in vertebrate evolution. Finally, the origin of a postotic skull and axial vertebrae appears to be associated with the origin of the gnathostomes.     

Cotyledon venation patterns in the Leguminosae: Mimosoideae

Cotyledon venation patterns are described for 131 species representing the four main tribes of the Mimosoideae. The range of variation in venation pattern falls within that described previously for the Caesalpinioideae and is consistent with the proposal that all patterns in the Leguminosae have been derived from a prototype with four vascular strands and a protoxylem trace in the petiole and seven primary veins in the lamina. Each tribe is characterized by a particular set of patterns, pattern frequencies and evolutionary trends. In the Mimoseae, correlations between venation pattern and cotyledon size and anatomy match those found in the Caesalpinioideae, but different correlations unique to these tribes occur in the Acacieae and Ingeae.     

Seed-coat anatomy of the non-pleurogrammic seeds in the tribe Ingeae (Leguminosae,Mimosoideae)

An anatomically complex structure of the seed-coat is a general characteristic of all members of the Leguminosae. Nonetheless, certain genera exhibit a particular type of seed, the “overgrown” seen that has been defined by a developmental criterion and by more simple anatomical features of the seed-coat. Only the last criterion seems suitable for unambiguously distinguishing this type of seed, which is here referred to as “overgrown-like.” This type appears to be apomorphic in the Ingeae (and probably in all the Leguminosae) and likely results from a heterochronic loss of tissue differentiation. Variations in this character may be useful at the generic level, and detailed anatomical observations reveal the occurrence of three distinct patterns. The high degree of correlation with other characters suggests that overgrown-like seeds have evolved separately at least three times in the Ingeae and thatPithecellobium s.s. may be polyphyletic. The overgrown-like seeds are likely to be an adaptive response to wet tropical climates.