Chemical Identity of a Rotting Animal-Like Odor Emitted from the Inflorescence of the Titan Arum (Amorphophallus titanum)

The titan arum, Amorphophallus titanum, is a flowering plant with the largest inflorescence in the world. The flower emits a unique rotting animal-like odor that attracts insects for pollination. To determine the chemical identity of this characteristic odor, we performed gas chromatography-mass spectrometry-olfactometry analysis of volatiles derived from the inflorescence. The main odorant causing the smell during the flower-opening phase was identified as dimethyl trisulfide, a compound with a sulfury odor that has been found to be emitted from some vegetables, microorganisms, and cancerous wounds.     

EVOLUTION OF REPRODUCTIVE STRATEGIES IN THE SEXUALLY DECEPTIVE ORCHID OPHRYS SPHEGODES: HOW DOES FLOWER‐SPECIFIC VARIATION OF ODOR SIGNALS INFLUENCE REPRODUCTIVE SUCCESS?

The orchid Ophrys sphegodes Miller is pollinated by sexually excited males of the solitary bee Andrena nigroaenea, which are lured to the flowers by visual cues and volatile semiochemicals. In O. sphegodes, visits by pollinators are rare. Because of this low frequency of pollination, one would expect the evolution of strategies that increase the chance that males will visit more than one flower on the same plant; this would increase the number of pollination events on a plant and therefore the number of seeds produced. Using gas chromatography-mass spectrometry (GC-MS) analyses, we identified more than 100 compounds in the odor bouquets of labellum extracts from O. sphegodes; 24 compounds were found to be biologically active in male olfactory receptors based on gas chromatography with electroantennographic detection (GC-EAD). Gas chromatography (GC) analyses of odors from individual flowers showed less intraspecific variation in the odor bouquets of the biologically active compounds as compared to nonactive compounds. This can be explained by a higher selective pressure on the pollinator-attracting communication signal. Furthermore, we found a characteristic variation in the GC-EAD active esters and aldehydes among flowers of different stem positions within an inflorescence and in the n-alkanes and n-alkenes among plants from different populations. In our behavioral field tests, we showed that male bees learn the odor bouquets of individual flowers during mating attempts and recognize them in later encounters. Bees thereby avoid trying to mate with flowers they have visited previously, but do not avoid other flowers either of a different or the same plant. By varying the relative proportions of saturated esters and aldehydes between flowers of different stem positions, we demonstrated that a plant may take advantage of the learning abilities of the pollinators and influence flower visitation behavior. Sixty-seven percent of the males that visited one flower in an inflorescence returned to visit a second flower of the same inflorescence. However, geitonogamy is prevented and the likelihood of cross-fertilization is enhanced by the time required for the pollinium deposited on the pollinator to complete its bending movement, which is necessary for pollination to occur. Cross-fertilization is furthermore enhanced by the high degree of odor variation between plants. This variation minimizes learned avoidance of the flowers and increases the likelihood that a given pollinator would visit several to many different plants within a population.     

THE POLLINATION BIOLOGY OF TUCKAHOE,PELTANDRA VIRGINICA (ARACEAE)

This study describes the specialized brood-site-based pollination system of Peltandra virginica Kunth (Araceae) with the chloropid fly Elachiptera formosa Loew and provides experimental evidence that gender-related changes in floral odor composition synchronize pollinator behavior with the blooming sequence. P. virginica is protogynous and does not self-pollinate because of a strong temporal separation in sexual function, and it is dependent upon insects for pollination because the spathe completely surrounds the spadix during the pistillate stage. Field observations conducted in central New Jersey and southeastern Pennsylvania showed that E. formosa is closely associated with P. virginica inflorescences. Within the floral chamber, E. formosa adults feed on pollen, mate, and find oviposition sites while the larvae complete their development. Although drosophilid and syrphid flies were collected and reared from inflorescences of both sexual stages, only E. formosa emerged from pistillate-stage inflorescences and adult E. formosa rapidly transferred fluorescent dye particles between inflorescences of both sexual stages in laboratory enclosures. These findings indicate that this fly species is the primary pollinator at our study locations. Field censuses demonstrated that although E. formosa visited P. virginica inflorescences of both sexual stages with equal frequency, the female flies preferentially oviposited within pistillate-stage inflorescences. Analysis of floral volatiles with gas chromatography-mass spectrometry showed that only the spathes emit the floral fragrance and that the composition of the floral volatiles changed during the transition from pistillate to staminate stage. A particularly noticeable change occurred in the emission ratio of the two primary floral odor components, 4,5,7-trimethyl-6,8-dioxabicyclo[3.2.1.]octane and an unidentified chemical analog with molecular weight of 142. The relationship between floral volatile composition and fly oviposition behavior was evaluated using sham inflorescences placed in the P. virginica study population. The sham inflorescences were constructed by covering spadices dissected from mature inflorescence buds with spathes excised from different-age inflorescences. After a 2-hr-long exposure period, sham inflorescences fitted with pistillate-stage spathes contained more E. formosa eggs than the sham inflorescences with staminate-stage spathes. However, the highest number of ovipositions occurred in sham inflorescences fitted with spathes excised from mature inflorescence buds. The fragrance emitted by these spathes was composed almost entirely of the two principal odor components. The decline in ovipositions observed in both real and sham inflorescences corresponded to an increase in the emission ratio of 4,5,7-trimethyl-6,8-dioxabicyclo[3.2.1.]octane to the molecular weight 142 analog. This suggests that gravid flies searching for oviposition sites used the ratio of the two primary floral volatile components as an inflorescence gender or age recognition cue.     

Role of EVERGREEN in the development of the cymose petunia inflorescence

Plants species diverge with regard to the time and place where they make flowers. Flowers can develop from apical meristems, lateral meristems, or both, resulting in three major inflorescence types known as racemes, cymes, and panicles, respectively. The mechanisms that determine a racemose architecture have been uncovered in Arabidopsis and Antirrhinum. To understand how cymes are specified, we studied mutations that alter the petunia inflorescence. Here we show that EVERGREEN (EVG) encodes a WOX homeodomain protein, which is exclusively expressed in incipient lateral inflorescence meristems (IMs), promoting their separation from the apical floral meristem (FM). This is essential for activation of DOUBLE TOP and specification of floral identity. Mutations that change the cymose petunia inflorescence into a solitary flower fully suppress the evg phenotype. Our data suggest a key role for EVG in the diversification of inflorescence architectures and reveal an unanticipated link between the proliferation and identity of meristems.     

Structure of the Cyperaceae Inflorescence

This work presents the basics for interpreting the adult inflorescence structure in Cyperaceae. It provides an analysis of the variations of the synflorescence and inflorescence structure in the family. Three types of synflorescence may be recognized in this family: a synflorescence with a foliate stem, a terminal inflorescence and a variable number of lateral inflorescences; a synflorescence with a foliate stem and only the terminal inflorescence; and a synflorescence with a scape and a terminal inflorescence. Variations in the structure and form of the inflorescences are related to variations in inflorescence branching, inflorescence homogenization degree, presence or absence of the distal part of the inflorescence, phyllotaxis, inflorescence position, types of bracts and leaves subtending branches, elongation of inflorescence internodes and spikelet structure. These variations are correlated with some of the developmental processes that give origin to the inflorescence.     

Organographic and ontogenetic studies on the inflorescence ofLespedeza cuneata (Dum.-Cours.) G. Don (Leguminosae)

Structure of inflorescence and its variation were organographically and ontogenetically studied inLespedeza cuneata (Dum.-Cours.) G. Don. An axillary inflorescence of the species forms a compound inflorescence which is composed of three or four component inflorescences. Each component inflorescence bears four (rarely six), three, two, or one flowers. Based on the arrangement of inflorescence phyllomes, the component inflorescence with four flowers is interpreted as a pseudoraceme bearing two shortened lateral shoots (partial inflorescences) each of which has two flowers. The component inflorescence with one flower appears to be terminated by the flower and to compose the cyme. Organographic observations revealed that the terminally located flower is not truly terminal, but axillary in origin. Ontogenetic observations showed that the apices of component inflorescence and partial inflorescence exist in early developmental stages in spite of variation in the form of component inflorescence. The terminally located flower in the cyme-like inflorescence was thus demonstrated to be laterally borne on the partial inflorescence axis. The component inflorescence composing the cyme-like one inL. cuneata is a reduced form in the number of partial inflorescences and of flowers from the pseudoraceme. The cyme-like inflorescence inL. cuneata resembles the inflorescence ofKummerowia.     

Hooks for mammal pollination?

Summary Two species of Banksia (family Proteaceae) studied in Australia were shown to be pollinated by small, non-flying mammals rather than by birds as previously thought, and to possess several adaptations appropriate for mammal-rather than bird-pollination: odor, troughs that channel excess nectar to the ground for attraction, open inflorescence structure for nectar accessibility, hooked wiry styles for effective pollen transfer, crepuscular and nocturnal nectar and pollen presentation, and copious nectar. This apparently is the first documentation with quantified data of pollination by non-flying mammals, although many other probable examples exist.     

Ontogeny and Organization of the Inflorescence in Balanophora

The locus of origin of the inflorescence in the tuber of BalanophoraJ.R. & G. Forst, is predetermined. A small nest of deep-seatedhypodermal cell-layers of the tuber, encircled by compositeconducting strands, becomes meristematic and organizes a morecompact club-shaped body. Its anterior broader end is sharplydelimited from the rest of the tuber tissue. One or two celllayers of the tuber abutting its broader end break down to forma lysigenous cavity that enlarges schizogenously. These changescoincide with cessation of growth of the composite conductingstrands. The inflorescence axis redifferentiates epidermal andhypodermal tissues prior to the initiation of flowers. The procambialsystem of the inflorescence originates from (1) the base ofthe inflorescence in the tuber and (2) the shoulder regionsof the composite conducting strands; the two extensively anastomosewith each other and yet remain wholly independent of the hostvasculature. Balanophora, inflorescence, ontogeny     

Morphological traffic between the inflorescence and the vegetative shoot in helobial monocotyledons

Previous studies of reproductive structures in the helobial monocotyledons (Alismatidae) indicate that partitioning between flower and inflorescence is not always clear (e.g.,Lilaea,Scheuchzeria) and that this may be the result of ancestral, unisexual modules coming together to form flowers and/or inflorescences. Later evolutionary changes may have included the inflorescence becoming involved or mixed in with vegetative growth. Substitution of vegetative buds for flowers is the simplest version, and there can be additional modifications to the growth behavior of the inflorescence, such as horizontal growth and dorsiventrality. In the Alismataceae and Limnocharitaceae the derivation of stolonlike structures from inflorescences is obvious: vegetative features have been incorporated into structures that are recognizably inflorescences. In the Hydrocharitaceae the interrelationships between the inflorescence and the vegetative body are much less well defined. We previously suggested forHydrocharis, where a single axillary complex can contain both inflorescence and stolons, that the stolon is basically a sterilized inflorescence and that features of the inflorescence have become incorporated into the vegetative body. Here we will explore this theme further for the Hydrocharitaceae, using information from within and outside the family.     

Development of the epiphyllous inflorescence of Phyllonoma integerrima (Turcz.) Loes.: implications for comparative morphology*

The inflorescence primordium of Phyllonoma integerrima (Turcz.) Loes. is initiated on the adaxial side of the leaf primordium. At about the same time, a vegetative bud is formed at the base of the same leaf primordium. The vascular anatomy is the same in the fertile and sterile leaves, except that in the fertile leaf an inflorescence trace departs from the midvein of the leaf at the point where the inflorescence is inserted. Neither the inception nor the procambial supply of the inflorescence provide evidence of “congenital fusion”of inflorescence and leaf. It is also argued that the idea of an “adventitious”origin of the inflorescence is not useful in this case. Consequences for our conception of shoot construction are pointed out. It is argued that positional changes in the initiation of organs is an evolutionary process that may have remarkable effects on plant construction.     

The inflorescence architecture of Petunia hybrida is modified by the Arabidopsis thaliana Ap2 gene.

We have introduced the Apetala2 (Ap2) gene of Arabidopsis thaliana into Petunia hybrida. Four out of 10 Ap2 transgenic plants flowered and exhibited an altered inflorescence architecture. Internode elongation suggests that the transition from the vegetative to the inflorescence phase does occur, although flower formation is delayed and the cymose branching pattern is not established. Instead, the inflorescence continues to produce bracts and eventually terminates in an aberrant flower with an excess of floral organs. New inflorescence branches then develop from the axillary meristems of the bracts, repeating the formation of a number of bracts before conversion into a terminal, aberrant flower. These results indicate that the Ap2 gene plays a role in the determination of inflorescence meristem identity, but not as a typical A-like function, adding to the existing doubt about the general role of Ap2 gene(s) in floral development.     

Testing the ontogenetic base for the transient model of inflorescence development

Backgrounds and Aims

Current research in plant science has concentrated on revealing ontogenetic processes of key attributes in plant evolution. One recently discussed model is the ‘transient model’ successful in explaining some types of inflorescence architectures based on two main principles: the decline of the so called ‘vegetativeness’ (veg) factor and the transient nature of apical meristems in developing inflorescences. This study examines whether both principles find a concrete ontogenetic correlate in inflorescence development.

Methods

To test the ontogenetic base of veg decline and the transient character of apical meristems the ontogeny of meristematic size in developing inflorescences was investigated under scanning electron microscopy. Early and late inflorescence meristems were measured and compared during inflorescence development in 13 eudicot species from 11 families.

Key Results

The initial size of the inflorescence meristem in closed inflorescences correlates with the number of nodes in the mature inflorescence. Conjunct compound inflorescences (panicles) show a constant decrease of meristematic size from early to late inflorescence meristems, while disjunct compound inflorescences present an enlargement by merging from early inflorescence meristems to late inflorescence meristems, implying a qualitative change of the apical meristems during ontogeny.

Conclusions

Partial confirmation was found for the transient model for inflorescence architecture in the ontogeny: the initial size of the apical meristem in closed inflorescences is consistent with the postulated veg decline mechanism regulating the size of the inflorescence. However, the observed biphasic kinetics of the development of the apical meristem in compound racemes offers the primary explanation for their disjunct morphology, contrary to the putative exclusive transient mechanism in lateral axes as expected by the model.     

Evolutionary origin of the Asteraceae capitulum: Insights from Calyceraceae

? Premise of the study: Phylogenies based on molecular data are revealing that generalizations about complex morphological structures often obscure variation and developmental patterns important for understanding the evolution of forms, as is the case for inflorescence morphology within the well-supported MGCA clade (Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae). While the basal families share a basic thyrsic/thyrsoid structure of their inflorescences, Asteraceae possesses a capitulum that is widely interpreted as a racemose, condensed inflorescence. Elucidating the poorly known inflorescence structure of Calyceraceae, sister to Asteraceae, should help clarify how the Asteraceae capitulum evolved from thyrsic/thyrsoid inflorescences. ? Methods: The early development and structure of the inflorescence of eight species (five genera) of Calyceraceae were studied by SEM, and patterns of evolutionary change were interpreted via phylogenetic character mapping. ? Key results: The basic inflorescence structure of Calyceraceae is a cephalioid (a very condensed botryoid/thyrsoid). Optimization of inflorescence characters on a DNA sequence-derived tree suggests that the Asteraceae capitulum derives from a simple cephalioid through two morphological changes: loss of the terminal flower and suppression of the cymose branching pattern in the peripheral branches. ? Conclusions: Widely understood as a condensed raceme, the Asteraceae capitulum is the evolutionary result of a very reduced, condensed thyrsoid. Starting from that point, evolution worked separately only on the racemose developmental control/pattern within Asteraceae and mainly on the cymose developmental control/pattern within Calyceraceae, producing head-like inflorescences in both groups but with very different diversification potential. We also discuss possible remnants of the ancestral cephalioid structure in some Asteraceae.     

Morphometric analysis of the genusHemerocallis L. (Liliaceae) in Korea

To better understand the patterns of variability and distributions ofHemerocallis in Korea, 53 locations were visited and measurements of 19 morphological and phenological characters were taken on plants directly from their natural habitats. For morphometric analysis, 10 plants from each of 34 populations and five herbarium specimens ofH. middendorffii were used and the data from 12 quantitative characters was analyzed using univariate analysis. Except the littoral populations of Cheju, Hong, Taehuksan, and Sohuksan Islands (H. hongdoensis M. Chung & S. Kang), three peninsular KoreanHemerocallis species can be recognized mainly in South Korea:H. hakuunensis Nakai (=H. micrantha Nakai, growing on southern, central, and northwestern Korea);H. thunbergii Baker (=H. coreana Nakai, found on southeastern and central Korea); andH. middendorffii Tr. et Mey. (central and northeastern Korea). Morphological and phenological features contributing to recognition of the three groups were; color of perianth, shape of roots, shape of inflorescence, flowering time, odor, length of inflorescence, width of the lowest bracts, length of perianth tube enclosing a ovary, width of the inner perianth lobes. Natural hybridization seems to be rare in KoreanHemerocallis. It appears that the KoreanHemerocallis species are relatively well characterized by their distribution patterns, phenology, and habitats compared with the JapaneseHemerocallis species.     

A novel gene Bractea (bra) controls the formation of an indeterminate bractless inflorescence in Arabidopsis thaliana

The morphological and genetic studies of the bra mutant of Arabidopsis thaliana (L.) Heynh. from the collection of the Department of Genetics and Breeding, Moscow State University, showed that the BRA gene controls the main stages of inflorescence development: it suppresses the development of leaflike organs subtending flowers (bracts) and inhibits the formation of the terminal flower. Inactivation of the BRA gene leads to the transition from the indeterminate bractless inflorescence characteristic of the family Cruciferaceae to the determinate bracteose inflorescence. The BRA gene plays a regulatory role and was probably involved in the conversion of the bracteose determinate inflorescence to the bractless indeterminate inflorescence during the origin of ancestral crucifers.     

Nonsteroidal moulting hormone agonists: effects on protein synthesis and cuticle formation in Colorado potato beetle larvae

Adult parasitoid wasps can learn to recognize specific resource-based cues, making them adept at locating essential resources within their native habitats. However, relatively little is known about their ability to recognize the odors emitted by flowers and extra-floral nectar glands. A novel test arena mimicking the distribution of nectaries within an umbelliferous inflorescence was developed to measure and compare the responses of two eulophid wasps, Edovum puttleri and Pediobius foveolatus, to nectar-based odors. Although both parasitoid species were able to associate nectar location with the odors emitted by both real nectaries (dill flowers and snap bean stipules) and artificial nectars (1 M sucrose solution scented with banana or lemon food flavoring), the responses of inexperienced wasps to nectary odors differed. While inexperienced E. puttleri displayed little attraction to the odor of either dill flowers or snap bean stipules, inexperienced P. foveolatus displayed a strong attraction to odors of both types of nectaries. However, once it had experienced foraging on either dill or snap bean nectar, E. puttleri responded to those nectary odors as strongly as did P. foveolatus. The responsiveness of both wasp species to the odor of artificial nectar greatly increased after they foraged on sugar solution scented with either banana or lemon odor. That parasitoid wasp species can differ in their ability to recognize food-based cues is of fundamental and applied importance.     

On the problems of the classification of Chinese bamboos with creeping rhizomes

So far, the 15 genera of bamboos with creeping rhizomes in China have been known. Based on the types and evolution of inflorescence, which have been discussed in the paper, and other criteria of classification, the opinions are given as follows: 1. Some species of Semiarundinaria spp. McClure and Sinobambusa spp. McClure should be separated and regarded as representing a new genus, because the racemes of the some species are different from false inflorescence (indeterminate infl.) of the type species of the two genera, and some other species should be transfered to the genus Pleioblastus. 2. On account of the similarities in the type of inflorescence and pistil, Brachystachyum may be combined with Semiarundinaria. 3. As the inflorescence, pistil and rhizome of Sinarundinaria are similar in those of Fargesia, the two genera should be combined. Due to the inflorescence of Fargesia is different from that of Thamnocalamus, therefore, these two genera must be kept separating. 4. Key to the Genera of Chinese bamboos with creeping rhizomes is provided inthis paper. But the key is available to flowering materials only, because the construction of it is mainly based on reproductive organs.     

Roles of odor, taste, and toxicity in the food preferences of lambs: implications for mimicry in plants

In the traditional sense, food ingestion consists of prehending, masticating, swallowing, and digesting plant matter. It is also possible to ingest plants without eating them. Volatile compounds are inhaled directly into the lungs and transported from the lungs into the bloodstream. Volatiles in high concentrations could presumably produce toxicosis, without an herbivore ever ingesting a plant in the customary sense. Volatile compounds may be aposematic, serving to warn potential predators of toxins in plants. We conducted three experiments to explore the roles of odor, taste, and toxicity in the food preferences of lambs. The first experiment determined if brief exposure to a novel odor followed by lithium chloride (LiCl)‐induced toxicosis caused lambs to avoid a familiar food that contained the odor. Lambs that sniffed coconut‐flavored barley and then received LiCl subsequently ate less coconut‐flavored barley than lambs that did not receive LiCl. The second experiment determined if lambs were deterred from eating a familiar food by the odor of Astragalus bisulcatus. A. bisulcatus is a malodorous (to humans) sulfur‐containing herb considered unpalatable and toxic. Neither odor nor intraruminal infusions of A. bisulcatus deterred lambs from feeding. The third experiment also determined how the degree of familiarity with the odor of A. bisulcatus, along with toxicosis, influenced preference of lambs for food with or without the odor of A. bisulcatus. Lambs with 8 d exposure to the odor but not given LiCl ate similar amounts of food, with and without the odor of A. bisulcatus, whereas lambs given LiCl showed a mild aversion to food with the odor during testing. Lambs with 1 d exposure to the odor but no LiCl ate similar amounts of food, with and without the odor, whereas lambs given LiCl showed a strong but transient aversion to food with the odor. Collectively, these findings show that lambs responded strongly to novel odors, but their response was transient and depended on the postingestive consequences of toxins and nutrients associated with odor inhalation. Thus, we submit that odor alone, in the absence of toxicosis or nociception, is not a deterrent to herbivores that continually sample foods and adjust intake based on the postingestive effects of toxins and nutrients. It also is unlikely that non‐toxic plants can mimic the odors of toxic plants to avoid herbivory (Batesian mimicry), unless the odors are indistinguishable by herbivores, again because herbivores constantly sample foods.     

The effects of herbivory and resource variability on the production of a second inflorescence by the desert lily, Pancratium sickenbergeri

We investigated the effects of herbivory by the dorcas gazelle, Gazella dorcas, on the production of a second inflorescence in a desert lily, Pancratium sickenbergeri (Amaryllidaceae), after the first inflorescence was eaten. In three populations exposed to different levels of herbivory, we conducted an inflorescence-clipping experiment in the flowering seasons of 1997, 1998 and 1999. The experiment was performed on inflorescences at both the emerging stage and the anthesis stage. Plants in each control group were undamaged. Clipped plants had a greater probability of producing a second inflorescence stalk than unclipped plants. In all the populations, the probability of producing a second inflorescence was greater when the first inflorescence was cut in the emerging stage than when it was cut in anthesis, indicating that there was a cost of compensation. In the population with the highest level of herbivory, the production of a second inflorescence stalk was related to resource availability in all 3 years. Plants produced as many flowers or fruits on the second stalk as on the first, indicating that potential reproductive output is double that usually achieved. Although we found no evidence for a trade-off between investment in first and second inflorescence stalks, the lower probability of producing a second inflorescence if heavy investment had been made in the first inflorescence indicates that compensation is costly. These results contradict the mutualism hypothesis that herbivory is beneficial to plants and provide support for the trade-off hypothesis that benefits of herbivory are proximal and are attained at an evolutionary cost.