Flower-like terminal structures in racemose inflorescences: a tool in morphogenetic and evolutionary research

Terminal flower-like structures (TFLS) occur in many angiosperms that possess indeterminate inflorescences such as spikes, racemes, or spadices. We describe and review TFLS in early-divergent angiosperms, especially the magnoliid order Piperales and the monocot order Alismatales, in which floral interpretation is controversial. Essentially similar TFLS occur in a wide range of taxa. Among magnoliids, they occur in some Piperales (Saururaceae and a few Piperaceae), but are absent from Chloranthaceae. Among monocots, they occur in some early-divergent families such as Acoraceae, Aponogetonaceae, Juncaginaceae, Potamogetonaceae, and Ruppiaceae. Similar TFLS with obscure organ identity are recorded in mutants of Arabidopsis. TFLS can often be interpreted as pseudanthia (close aggregations of reduced flowers), but in some cases the entire terminal pseudanthium is very similar to a true flower. In some cases, elaborated TFLS could therefore have given rise to what are normally termed 'true' (i.e. euanthial) flowers. Data presented here on terminal pseudanthia in Potamogeton and Ruppia support a pseudanthial evolutionary origin of reproductive units in the alismatid families Zannichelliaceae and Cymodoceaceae. Furthermore, in some alismatid species, either the entire inflorescence apex or an individual primordium at or near the inflorescence tip can be transformed into a filamentous or tubular (or intermediate) structure. A tubular structure enclosing stamens and carpels is described in Piper. This indicates that pseudanthium formation can provoke morphological novelties, perhaps due to new patterns of overlap between expression zones of regulatory genes and/or new spatial constraints.     

Comparative ontogeny of the cyathium in Euphorbia (Euphorbiaceae) and its allies: exploring the organ-flower-inflorescence boundary

We present a detailed comparative ontogenetic analysis of pseudanthia of representatives of all three subtribes of Euphorbieae (Euphorbiinae, Neoguillauminiinae, Anthosteminae) in order to clarify their homologies and interpretation. The cyathium of Euphorbia and its allies (subtribe Euphorbiinae) closely resembles a bisexual flower but is traditionally interpreted as an inflorescence bearing clusters of highly reduced male flowers surrounding a single terminal female flower. Previously unreported characters are (1) male flowers formed one above the other in the male inflorescences of some Euphorbiinae, (2) late-developing perianthlike structures in some male flowers of Neoguillauminia cleopatra, (3) evidence for a bracteate origin of the female perianth in Anthosteminae and Neoguillauminiinae, and (4) spatiotemporally independent formation of abscission zone and perianth. Indistinct boundaries between inflorescence, flower, and floral organs demonstrate that defining the cyathium neither as an inflorescence nor as a flower is entirely satisfactory and indicate a "hybrid" flower/inflorescence nature of the cyathium. Based on our current knowledge and the existing phylogenetic context, it is most parsimonious to suggest that the cyathium evolved from a determinate thyrse with a terminal female flower surrounded by dichasial male partial inflorescences. We speculate that the cyathium was formed because of strong condensation and possible overlap between expression zones of regulatory genes.     

Enhanced inflorescence development in bougainvillea "san diego red" by removal of young leaves and cytokinin treatments

Removal of young leaves and application of the cytokinin, N-benzyla-α-(tetrahydro-2H-pyran-2yl)-adenine promote inflorescence development in Bougainvillea “San Diego Red”. Defoliation greatly increased the amount of assimilate accumulated at the shoot tip 1 to 2 days after treatment. Cytokinin applications further increased the amount accumulated and this increase was apparent 4 days before morphological changes could be detected at the inflorescence axes. Short days promoted inflorescence development and also increased assimilate accumulation at the reproductive axes; thus, it is suggested that the role of short day induction in bougainvillea may be that of redirecting the flow of assimilates, perhaps by its influence on cytokinin synthesis and distribution.     

Gravitropic response of inflorescence stems in Arabidopsis thaliana.

We have characterized the gravitropic response of inflorescence stems in Arabidopsis thaliana. When the inflorescence stems were placed horizontally, they curved upward about 90 degrees within 90 min in darkness at 23 degrees C, exhibiting strong negative gravitropism. Decapitated stem segments (without all flowers, flower buds, and apical apices) also showed gravitropic responses when they included the elongation zone. This result indicates that the minimum elements needed for the gravitropic response exist in the decapitated inflorescence stem segments. At least the 3-min gravistimulation time was sufficient to induce the initial curvature at 23 degrees C after a lag time of about 30 min. In the gravitropic response of inflorescence stems, (a) the gravity perception site exists through the elongating zone, (b) auxin is involved in this response, (c) the gravitropic curvature was inhibited at 4 degrees C but at least the gravity perception step could occur, and (d) two curvatures could be induced in sequence at 23 degrees C by two opposite directional horizontal gravistimulations at 4 degrees C.     

CYCLOIDEA 2 Clade Genes: Key Players in the Control of Floral Symmetry,Inflorescence Architecture,and Reproductive Organ Development

Undoubted lines of evidence point out that members of CYCLOIDEA (CYC) 2 clade are essential players to control flower symmetry and, amusingly, also are determinants of capitula architecture (pseudanthium). In several species, CYC-like genes influence the androecium patterning, but to date, the function of these genes in the development of gynoecium organs is less clear. In this review, we first reported details about floral symmetry and an overview of genes and molecular mechanisms regulating the development of zygomorphism in different angiosperm lineages (e.g., basal and core eudicots and monocots). Then, we paid emphasis on the role of CYC-like genes in the development of heterogamous inflorescence of sunflower as well as other Asteraceae and some species within the Dipsacaceae family. Helianthus annuus is particularly attractive because it represents a useful model to study the role of CYC-like genes on shaping floral corolla as well as the differentiation of reproductive organs in different flowers of pseudanthia. A special attention was reserved to inflorescence morphology mutants of sunflower (i.e., Chrysanthemoids2 and tubular ray flower) because they provide useful information on the role of CYC-like genes in the radiate capitulum evolution. Finally, we discuss data from literature to suggest that CYC-like genes are also co-opted to regulate stamen and carpel differentiation likely throughout their interaction with the cell cycle and flower organ identity genes. The recruitment of reproductive organs in ray flowers also supports the phylogenetic origin of a radiate inflorescence of sunflower from a discoid capitulum and suggests that in sterile zygomorphic ray flower primordia the latent identity to differentiate both microsporangium and macrosporangium was conserved.     

Nonflowers near the base of extant angiosperms? Spatiotemporal arrangement of organs in reproductive units of Hydatellaceae and its bearing on the origin of the flower

Reproductive units (RUs) of Trithuria, the sole genus of the early-divergent angiosperm family Hydatellaceae, are compared with flowers of their close relatives in Cabombaceae (Nymphaeales). Trithuria RUs combine features of flowers and inflorescences. They differ from typical flowers in possessing an "inside-out" morphology, with carpels surrounding stamens; furthermore, carpels develop centrifugally, in contrast to centripetal or simultaneous development in typical flowers. Trithuria RUs could be interpreted as pseudanthia of two or more cymose partial inflorescences enclosed within an involucre, but the bractlike involucral phyllomes do not subtend partial inflorescences and hence collectively resemble a typical perianth. Teratological forms of T. submersa indicate a tendency to fasciation and demonstrate that the inside-out structure-the primary feature that separates RUs of Hydatellaceae from more orthodox angiosperm flowers-can be at least partially modified, thus producing a morphology that is closer to an orthodox flower. The Trithuria RU could be described as a "nonflower", i.e., a structure that contains typical angiosperm carpels and stamens but does not allow recognition of a typical angiosperm flower. The term nonflower could combine cases of secondary loss of flower identity and cases of a prefloral condition, similar to those that gave rise to the angiosperm flower. Nonhomology among some angiosperm flowers could be due to iterative shifts between nonfloral construction and flower/inflorescence organization of reproductive organs. Potential testing of these hypotheses using evolutionary-developmental genetics is explored using preliminary data from immunolocalization of the floral meristem identity gene LEAFY in T. submersa, which indicated protein expression at different hierarchical levels.     

AtHVA22 gene family in Arabidopsis: phylogenetic relationship,ABA and stress regulation,and tissue-specific expression

HVA22 is an ABA- and stress-inducible gene first isolated from barley (Hordeum vulgare L.). Homologues of HVA22 have been found in plants, animals, fungi and protozoa, but not in prokaryotes, suggesting that HVA22 plays a unique role in eukaryotes. Five HVA22 homologues, designated AtHVA22a, b, c, d and e, have been identified in Arabidopsis. These five AtHVA22 homologues can be separated into two subfamilies, with AtHVA22a, b and c grouped in one subfamily and AtHVA22d and e in the other. Phylogenetic analyses show that AtHVA22d and e are closer to barley HVA22 than to AtHVA22a, b and c, suggesting that the two subfamilies had diverged before the divergence of monocots and dicots. The distribution and size of exons of AtHVA22 homologues and barley HVA22 are similar, suggesting that these genes are descendents of a common ancestor. AtHVA22 homologues are differentially regulated by ABA, cold, dehydration and salt stresses. These four treatments enhance AtHVA22a, d and e expression, but have little or even suppressive effect on AtHVA22c expression. ABA and salt stress induce AtHVA22b expression, but cold stress suppresses ABA induction of this gene. Expression of AtHVA22d is the most tightly regulated by these four treatments among the five homologues. In general, AtHVA22 homologues are expressed at a higher level in flower buds and inflorescence stems than in rosette and cauline leaves. The expression level of these homologues in immature siliques is the lowest among all tissues analyzed. It is suggested that some of these AtHVA22 family members may play a role in stress tolerance, and others are involved in plant reproductive development.     

Influence of growth regulators on somatic embryogenesis,plantlet regeneration,and post-transplant survival of Echinochloa frumentacea

After placement on Murashige and Skoog's basal medium supplemented with 3–5 mg/l 2,4-D, immature inflorescence expiants of Echinochloa frumentacea gave rise to three distinct types of callus: a) loosely arranged and soft; b) compact and translucent; c) compact, sticky and mucilaginous. Somatic embryo formation occurred in type b callus in about 18–24 d. Callus types a and c did not produce somatic embryos. The highest percentage of cultures exhibiting somatic embryogenesis occurred on the medium containing 5 mg/l 2,4-D and 0.5 mg/l kinetin. Somatic embryos also formed directly on the inflorescence (without intervening callus formation) in about 15% of the expiants placed on this medium. The addition of paclobutrazol or uniconazole (0.25 or 1 mg/l) to the medium had no influence on the percentage of cultures exhibiting direct somatic embryogenesis, but paclobutrazol slightly increased the mean number of somatic embryos per culture. Many of the callus-derived somatic embryos germinated when subcultured on basal MS medium supplemented with kinetin. Addition of paclobutrazol or uniconazole to the culture medium at 0.25 or 1 mg/l decreased somatic embryo germination and shoot elongation but increased root length and leaf width. Both paclobutrazol and uniconazole increased survival of the plantlets following transplanting to soil. Increased post-transplant survival was accompanied by reduced water loss from plantlets produced on culture media containing triazoles.     

Cytokinin-induced <Emphasis Type="Italic">VvTFL1A</Emphasis> expression may be involved in the control of grapevine fruitfulness

Grapevine bud fruitfulness is determined by the differentiation of uncommitted meristem (UCM) into either tendril or inflorescence. Since tendril and inflorescence differentiation have long been considered sequential steps in inflorescence development, factors that control the progression of floral meristem development may regulate the final outcome of UCM differentiation, and thus affect fruitfulness. A comparison of the expression profiles of the master regulators of floral meristem identity (FMI) during development of fruitful and non-fruitful buds along the same cane allowed associating the expression of a homolog of terminal flower 1 (TFL1, a negative regulator of FMI) to fruitful buds, and the expression of positive FMI regulators to non-fruitful buds. Combined with (a) cytokinin-induced upregulation of VvTFL1A expression in cultured tendrils, which accompanied cytokinin-derived tendril transformation into branched, inflorescence-like structures, (b) positive regulation of VvTFL1A expression by cytokinin, which was demonstrated in transgenic embryonic culture expressing GUS reporter under the control of VvTFL1A promoter, and (c) a significantly higher level of active cytokinins in fruitful positions, the data may support the assumption of cytokinin-regulated VvTFL1A activity’s involvement in the control of inflorescence development. Such activity may delay acquisition of FMI and allow an extended branching period for the UCM, resulting in the differentiation of inflorescence primordia.     

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.     

The movement patterns of carpenter beesXylocopa micans and bumblebeesBombus pennsylvanicus onPontederia cordata inflorescences

The movement patterns of carpenter bees (Xylocopa micans) and bumblebees (Bombus pennsylvanicus) foraging for nectar on vertical inflorescences ofPontederia cordata were studied near Miami, Florida. The floral biology ofP. cordata is unique in several ways: (a) many short-lived flowers per inflorescence, (b) constant nectar production throughout the life span of each flower, and (c) abscence of vertical patterning of nectar and age of flowers. Inflorescences ranged between 3.5 and 15.8 cm long and had between 9 and 55 open flowers. Both carpenter bees and bumblebees arrived mostly on the bottom third of the inflorescence and left after visiting flowers on the top third of the inflorescence. The departure position from the inflorescence was higher up than observed in studies of other insect pollinators foraging on other speces of plants. This pattern of departure probably occurs in the absence of a vertical gradient of nectar or floral morphology.     

Morphology and anatomy of inflorescence and inflorescence axis in Paepalanthus sect. Diphyomene Ruhland (Eriocaulaceae,Poales) and its taxonomic implications

Paepalanthus sect. Diphyomene has inflorescences arranged in umbels. The underlying bauplan seems however to be more complex and composed of several distinct subunits. Despite appearing superficially very similar, the morphology and anatomy of the inflorescences can supply useful information for the understanding of the phylogeny and taxonomy of the group. Inflorescences of Paepalanthus erectifolius, Paepalanthus flaccidus, Paepalanthus giganteus, and Paepalanthus polycladus were analyzed in regard to branching pattern and anatomy. In P. erectifolius, P. giganteus and P. polycladus the structure is a tribotryum, with terminal dibotryum, and with pherophylls bearing lateral dibotrya. In P. flaccidus, the inflorescence is a pleiobotryum, with terminal subunit, and without pherophylls. Secondary inflorescences may occur in all species without regular pattern. Especially when grown in sites without a pronounced seasonality, the distinction between enrichment zone (part of the same inflorescence) and new inflorescences may be obscured. The main anatomical features supplying diagnostic and phylogenetic information are as follows: (a) in the elongated axis, the thickness of the epidermal cell walls and the cortex size; (b) in the bracts, the quantity of parenchyma cells (c) in the scapes, the shape and the presence of a pith tissue. Therefore, P. sect. Diphyomene can be divided in two groups; group A is represented by P. erectifolius, P. giganteus and P. polycladus, and group B is represented by P. flaccidus. The differentiation is based in both, inflorescence structure and anatomy. Group A presents a life cycle and anatomical features similar to species of Actinocephalus. Molecular trees also point that these two groups are closely related. However, inflorescence morphology and blooming sequence are different. Species of group B present an inflorescence structure and anatomical features shared with many genera and species in Eriocaulaceae. The available molecular and morphology based phylogenies still do not allow a precise allocation of the group in the bulk of basal species of Paepalanthus collocated in P. sect. Variabiles. The characters described and used here supply however important information towards this goal.     

Developmental Changes of TGF-β1 and Smads Signaling Pathway in Intestinal Adaption of Weaned Pigs

Weaning stress caused marked changes in intestinal structure and function. Transforming growth factor-β1 (TGF-β1) and canonical Smads signaling pathway are suspected to play an important regulatory role in post-weaning adaptation of the small intestine. In the present study, the intestinal morphology and permeability, developmental expressions of tight junction proteins and TGF-β1 in the intestine of piglets during the 2 weeks after weaning were assessed. The expressions of TGF-β receptor I/II (TβRI, TβRII), smad2/3, smad4 and smad7 were determined to investigate whether canonical smads signaling pathways were involved in early weaning adaption process. The results showed that a shorter villus and deeper crypt were observed on d 3 and d 7 postweaning and intestinal morphology recovered to preweaning values on d 14 postweaning. Early weaning increased (P<0.05) plasma level of diamine oxidase (DAO) and decreased DAO activities (P<0.05) in intestinal mucosa on d 3 and d 7 post-weaning. Compared with the pre-weaning stage (d 0), tight junction proteins level of occludin and claudin-1 were reduced (P<0.05) on d 3, 7 and 14 post-weaning, and ZO-1 protein was reduced (P<0.05) on d 3 and d 7 post-weaning. An increase (P<0.05) of TGF-β1 in intestinal mucosa was observed on d 3 and d 7 and then level down on d 14 post-weaning. Although there was an increase (P<0.05) of TβR II protein expression in the intestinal mucosa on d3 and d 7, no significant increase of mRNA of TβRI, TβRII, smad2/3, smad4 and smad7 was observed during postweaning. The results indicated that TGF-β1 was associated with the restoration of intestinal morphology and barrier function following weaning stress. The increased intestinal endogenous TGF-β1 didn''t activate the canonical Smads signaling pathway.     

Flower-like heads from flower-like meristems: pseudanthium development in <Emphasis Type="Italic">Davidia involucrata</Emphasis> (Nyssaceae)

Flower-like inflorescences (pseudanthia) have fascinated botanists for a long time. They are explained as condensed inflorescences implying that the pseudanthium develops from an inflorescence meristem (IM). However, recent developmental studies identified a new form of reproductive meristem, the floral unit meristem (FUM). It differs from IMs by lacking acropetal growth and shares fractionation, expansion and autonomous space filling with flower meristems (FM). The similarity among FUMs and FMs raises the question how far flower-like heads originate from flower-like meristems. In the present paper, pseudanthium development in Davidia involucrata is investigated using scanning electron microscopy. D. involucrata has pincushion-shaped heads composed of densely aggregated, perianthless flowers and associated with two large showy bracts. Early developmental stages show a huge naked FUM. The FMs appear almost simultaneously and lack subtending bracts. With ongoing FUM expansion new space is generated which is immediately used by further FM fractionation. The heads have only staminate flowers or are andromonoecious with staminate and a single perfect flower in oblique position. All FMs lack perianth structures and fractionate a variable number of stamen primordia. The perfect FM is much larger than the staminate FMs and forms a syncarpous gynoecium with inferior ovary. Pseudanthium development in D. involucrata confirms the morphogenetic similarity to FMs as to acropetal growth limitation, meristem expansion and fractionation. It thus should not be interpreted as a condensed inflorescence, but as a flower equivalent. Furthermore as the FUM develops inside a bud, its development is considered to be influenced by mechanical pressure. The oblique position of the perfect flower, the developmental delay of the proximal flowers, and the variable number of stamens which were observed in the pseudanthium development, can be caused by mechanical pressure. Next to the Asteraceae, D. involucrata offers a further example of a pseudanthium originating from a FUM. More knowledge on FUMs is still needed to understand diversification and evolution of flower-like inflorescences.     

New concepts and methods of standardizing predictive value, accuracy and incorrect diagnostic rate

OBJECTIVE: To express the value of a diagnostic test under standardized and comparable conditions. STUDY DESIGN: Four new concepts of standardizing positive predictive value (SPPV), standardizing negative predictive value (SNPV), standardizing accuracy (SAc) and standardizing an incorrect diagnostic test were developed. The theoretical positive predictive value (SPPV), theoretical negative predictive value (SNPV), theoretical accuracy (SAc) and theoretical incorrect diagnosis rate (SIDR), which are not affected by a different constituent ratio of disease and nondisease groups and are obtained under the theoretical standard condition that the sample size in the disease group equals that in the nondisease group, were defined. Based on these concepts and the principles and methods of statistics and evaluation of diagnostic tests, corresponding formulas were deduced. RESULTS: The formulas are: SPPV = a(b + d)/[a(b + d) + b(a + c)] = Se/(1 + Se - Sp), SNPV = d(a + c)/[c(b + d) + d(a + c)] = Sp/(1 - Se + Sp), SAc = [a(b + d) + d(a + c)]/[2(a + c)(b + d)] = (Se + Sp)/2, and SIDR = [b(a + c) + c(b + d)]/[2(a + c)(b + d)] = (2 - Se - Sp)/2. Here, a, b, c and d refer to the case numbers of true positives, false positives, false negatives and true negatives; Se and Sp refer, respectively, to sensitivity and specificity. CONCLUSION: SPPV, SNPV, SAc and SIDR are very useful for expressing and evaluating the value of a diagnostic test under standardized and comparable conditions.     

Evolutionary and morphometric implications of morphological variation among flowers within an inflorescence: a case-study using European orchids

BACKGROUND AND AIMS: This study explores the previously largely ignored morphological variation that occurs among flowers within a single inflorescence. METHODS: Variation in four metric parameters (labellum length and width, spur length and width) that together strongly influence pollination frequency is documented within the simple racemose inflorescences of eight individuals that represent a primary hybrid and six species of European orchids. KEY RESULTS: Regression of each parameter against the location of each flower on the inflorescence, and calculation of correlation coefficients for each pair of parameters within each inflorescence, demonstrate significant decoupling of labellum and spur development, despite the fact that they are different portions of the same floral organ. Spur length and diameter are constant across inflorescences of Dactylorhiza other than the vestigial-spurred D. viridis, whereas in other genera spur length declines in parallel with labellum dimensions. These differences are likely to reflect selection pressures or developmental constraints. Strong negative deviations from the regression line for one or more parameters are evident in occasional flowers, occurring most frequently in the lowermost and uppermost one or two flowers, and so reflecting transitions in meristematic behaviour. Thus, population-level morphometric studies are best conducted on flowers taken from approximately the mid-point of the inflorescence. Moreover, in the two relatively large inflorescences where lower flowers were removed for measurement before the upper flowers had opened, labellum size increased significantly in the flowers immediately above the excisions, suggesting that excision liberated resources that were diverted into the opening buds. Repeat measurement of all flowers from one selected inflorescence demonstrated typical measurement errors of only +/- 30-80 micro m, irrespective of the size of the structure studied. If flowers are not mounted and measured immediately following excision, modest negative deviations of 30-50 micro m result from post-mounting shrinkage; this occurs less rapidly in the spur than in the thinner labellum, which should therefore be measured first. Variation in all four parameters among all the flowers of a single inflorescence is between 42 % and 107 % of that observed between a similar number of flowers sampled from a consistent location on different (but conspecific and coexisting) inflorescences. CONCLUSIONS: This result demonstrates the strong influence of epigenesis on flower morphology and further emphasizes the importance of (a) sampling from a consistent location within the inflorescences under comparison, (b) interpreting morphometric ordinations hierarchically, building from individuals to infraspecific taxa and species via populations, and (c) considering in any microevolutionary study the potentially profound effects of the cline in flower size within each inflorescence.     

Adaptation of Grapevine Flowers to Cold Involves Different Mechanisms Depending on Stress Intensity

Grapevine flower development and fruit set are influenced by cold nights in the vineyard. To investigate the impact of cold stress on carbon metabolism in the inflorescence, we exposed the inflorescences of fruiting cuttings to chilling and freezing temperatures overnight and measured fluctuations in photosynthesis and sugar content. Whatever the temperature, after the stress treatment photosynthesis was modified in the inflorescence, but the nature of the alteration depended on the intensity of the cold stress. At 4°C, photosynthesis in the inflorescence was impaired through non-stomatal limitations, whereas at 0°C it was affected through stomatal limitations. A freezing night (−3°C) severely deregulated photosynthesis in the inflorescence, acting primarily on photosystem II. Cold nights also induced accumulation of sugars. Soluble carbohydrates increased in inflorescences exposed to −3°C, 0°C and 4°C, but starch accumulated only in inflorescences of plants treated at 0 and −3°C. These results suggest that inflorescences are able to cope with cold temperatures by adapting their carbohydrate metabolism using mechanisms that are differentially induced according to stress intensity.     

Heat Production in the Voodoo Lily (Sauromatum guttatum) as Monitored by Infrared Thermography

The pattern of surface temperatures of the inflorescence of Sauromatum guttatum was investigated by using an infrared camera. The male flowers are weakly thermogenic on the first day of inflorescence opening (D-day) as well as on the next day (D + 1), reaching 0.5 to 1°C above ambient temperature. The appendix (the upper sterile part of the inflorescence) is highly thermogenic on D-day, reaching 32°C, and is faintly thermogenic on D + 1, reaching 1°C above ambient temperature. The lower part of the spadix, close to the female flowers, is also thermogenic on D-day and D + 1, reaching a temperature similar to that of the appendix only on D + 1. Salicylic acid does not induce heat production in the lower part of the spadix, as it does in the appendix. Respiration of tissue slices obtained from the appendix shows that the capacity for cyanide-insensitive respiration is present in young and mature appendices. This alternative respiratory pathway is not, however, utilized in young appendix tissue, but is engaged during the maturation of that tissue.