Domestication of cardamom (Elettaria cardamomum) in Western Ghats, India: divergence in productive traits and a shift in major pollinators

Background and Aims

Elettaria cardamomum, a highly priced spice, is native to the Western Ghats of South India. Wild populations still occur in isolated patches in their natural habitats; however, much of today''s commercial product comes from cultivated sources. There is no information on domestication-related traits of this species; the main objective of this study was to compare wild and cultivated populations of cardamom in terms of vegetative and reproductive features in order to identify domestication syndromes and to examine whether the two populations have developed reproductive barriers.

Methods

Two wild populations and five cultivated plantations were used for the present study. Vegetative and floral traits, flowering phenology, pollination biology and breeding systems of wild and cultivated populations were compared. Effective pollinators amongst floral visitors were identified by confirming pollen transfer as well as by fruit set following their visit to virgin flowers. Manual pollinations were carried out in order to study the breeding systems of the two populations and reproductive barriers, if any, between them.

Key Results

Several productive traits including the number of branches, number of inflorescences, and total number of flowers per clump, number of flowers that open each day, the duration of flowering, the length of the flower and the amount of nectar per flower are significantly greater in cultivated cardamom. The principal pollinators in wild cardamom are solitary bees, Megachile sp. and two species of Amegilla, whereas those in cultivated cardamom are the social bees Apis dorsata, A. cerana and Trigona iridipennis. Both the wild and cultivated populations are self-compatible and there are no reproductive barriers between the two populations.

Conclusions

Domestication in cardamom has brought about significant changes in vegetative and reproductive traits and a shift in effective pollinators from native solitary bees to social bees. The shift in pollinators seems to be due to the availability of a large number of flowers for prolonged periods in cultivated cardamom that can attract and sustain social bees, rather than due to co-evolution of the flower and the pollinator.Key words: Elettaria cardamomum, wild cardamom, domestication, Amegilla sp., Apis cerana, Apis dorsata, Megachile sp., pollination efficiency, solitary bees, social bees     

月季花瓣数量遗传分析

以‘窄叶藤本月季花’(Rosa chinensis‘Zhaiye Tengben Yuejihua’)ב月月粉’(R.chinensis‘Old Blush’)杂交群体为材料,分析其花瓣数量的分离特点,对单瓣花与重瓣花的花芽分化过程进行观察,并对花瓣、雄蕊及瓣化雄蕊进行表皮细胞超微结构的观察。结果显示:杂交群体的花瓣数量分离明显,出现从5~54片的连续变异;花瓣数量、瓣化雄蕊数量、雌蕊数量的遗传模型为2MG-AD(2对加性-显性-上位性主基因控制),雄蕊数量的遗传模型为0MG(无主基因控制);月季重瓣花形成的原因为雄蕊瓣化,重瓣花形成的关键时期为雄蕊原基形成后期,可见到雄蕊瓣化为花瓣的现象;月季瓣化雄蕊的表皮细胞形态、褶皱程度介于单瓣花花瓣和重瓣花外轮花瓣之间。     

Evolution of petal epidermal micromorphology in Leguminosae and its use as a marker of petal identity

Background and Aims

The legume flower is highly variable in symmetry and differentiation of petal types. Most papilionoid flowers are zygomorphic with three types of petals: one dorsal, two lateral and two ventral petals. Mimosoids have radial flowers with reduced petals while caesalpinioids display a range from strongly zygomorphic to nearly radial symmetry. The aims are to characterize the petal micromorphology relative to flower morphology and evolution within the family and assess its use as a marker of petal identity (whether dorsal, lateral or ventral) as determined by the expression of developmental genes.

Methods

Petals were analysed using the scanning electron microscope and light microscope. A total of 175 species were studied representing 26 tribes and 89 genera in all three subfamilies of the Leguminosae.

Key Results

The papilionoids have the highest degree of variation of epidermal types along the dorsiventral axis within the flower. In Loteae and genistoids, in particular, it is common for each petal type to have a different major epidermal micromorphology. Papillose conical cells are mainly found on dorsal and lateral petals. Tabular rugose cells are mainly found on lateral petals and tabular flat cells are found only in ventral petals. Caesalpinioids lack strong micromorphological variation along this axis and usually have only a single major epidermal type within a flower, although the type maybe either tabular rugose cells, papillose conical cells or papillose knobby rugose cells, depending on the species.

Conclusions

Strong micromorphological variation between different petals in the flower is exclusive to the subfamily Papilionoideae. Both major and minor epidermal types can be used as micromorphological markers of petal identity, at least in papilionoids, and they are important characters of flower evolution in the whole family. The molecular developmental pathway between specific epidermal micromorphology and the expression of petal identity genes has yet to be established.Key words: Epidermis, Fabaceae, Papilionoideae, Caesalpinioideae, Mimosoideae, petal surface, scanning electron microscopy, papillose conical cells, tabular rugose cells, tabular flat cells, organ identity     

Petal Abscission in Rose Flowers: Effects of Water Potential, Light Intensity and Light Quality

Petal abscission was studied in roses (Rosa hybrida L.), cvs.Korflapei (trade name Frisco), Sweet Promise (Sonia) and CaraMia (trade name as officially registered cultivar name). Unlikeflowers on plants in greenhouses, cut flowers placed in waterin the greenhouse produced visible symptoms of water stress,depending on the weather during the experiment and on the cultivar.Cut Frisco roses showed no visible signs of water stress andthe time to petal abscission was as in uncut flowers. In Soniaroses the symptoms of water stress varied from mild to severe,and the number of flowers in which the petals abscised variedfrom 100% (mild stress) to 0% (severe stress). An antimicrobialcompound in the vase water of Sonia roses, or removal of theleaves, alleviated the symptoms of water stress and increasedthe number of stems in which the petals abscised. Cut Cara Miaroses showed severe symptoms of water stress in all experimentsand petal abscission was found in only a few flowers, even whenthe stems were placed at 20 °C and low photon flux (15 µmolm^-2s^-1). Abscission in Sonia and Cara Mia roses was low or absentwhen the water potential of the leaves reached values below-2.0 MPa within the first 5 d of the experiment; such low valueswere not reached in Frisco roses. Addition of sucrose to the vase solution, together with an effectiveantimicrobial compound, had no effect on the time to petal abscission,at any light intensity. Placing flowers in far-red light alsohad no effect on abscission, compared with flowers placed inred light or white light of the same photon fluence. It is concluded that petal abscission in the rose cultivarsstudied is not affected by their water status unless the plantsreach a low water potential (about -2 MPa) early on during vaselife. Petal abscission is not inhibited by low light intensitynor affected by the Pr/Pfr ratio. Abscission; light intensity; petals; phytochrome; Rosa hybrida L.; rose; sugars; water potential     

Floral characteristics affect susceptibility of hybrid tea roses, Rosa x hybrida, to Japanese beetles (Coleoptera: Scarabaeidae)

The Japanese beetle, Popillia japonica Newman, feeds on the flowers and foliage of roses. Rosa x hybrida. Beetles attracted to roses land almost exclusively on the flowers. This study evaluated characteristics of rose flowers including color, size, petal count and fragrance, as well as height of plants and blooms within plant as factors in attractiveness to Japanese beetles. Artificial flowers that had been painted to match the spectral reflectance of real blooms were attached to potted nonflowering rose plants in the field and the number of beetles that landed on each model was recorded. More beetles landed on the yellow- and white-colored flower models than on the five other bloom colors that were tested. Large (15 cm diameter) yellow flower models attracted more beetles than did smaller (8 cm diameter) yellow models. There was no difference in beetle response to yellow flower models of the same size that differed in bloom complexity (i.e., number of petals). Experiments in which blooming rose plants were elevated above controls, or in which flower models were placed at different heights within plant canopies, failed to support the hypothesis that height per se accounts for beetles' attraction to flowers over leaves. Attractiveness of selected rose cultivars that varied in fragrance and flower color also was evaluated in the field. Yellow-flowered cultivars were more susceptible than those with red flowers, regardless of fragrance intensity as rated by breeders. Growing cultivars of roses that have relatively dark and small-sized blooms may have some benefit in reducing Japanese beetles' attraction to roses.     

The Abscission of Rose Petals

Petal abscission was studied in twelve hybrid tea rose (Rosahybrida L.) cultivars. At about 20 °C the time to petalabscission in uncut stems in greenhouses was the same as incut stems placed in water in the greenhouse or in a climate-controlledroom. The time between petal unfolding and abscission dependedon the cultivar, and varied between 12 and 35 d. The time topetal abscission of the cultivars was inversely correlated withtheir flower diameter at full bloom (linear regression, r² =0·82). In the cultivars with a relatively large flowerdiameter (10-18 cm) the petals fell without visible desiccationsymptoms, whereas in the group with a small diameter the petalswere partially or fully desiccated when shed. Fertilization occurred in some flowers of a few cultivars studied.In cultivars with a relatively large flower diameter (Papa Meilland,Cocktail, Dr. Verhage, Tineke) it had no effect on the timeto abscission in Motrea, Europa, and Carolien roses, which bearsmall flowers, the petals fell after fertilization, whereasin unfertilized flowers of the latter group of cultivars anabscission zone just above the uppermost node became activeand all parts above this node (pedicel and flower) turned brownand desiccated, though remained attached for more than a month. It is concluded that in the cultivars investigated: (a) thetime to petal abscission was inversely related to their flowerdiameter, (b) abscised petals were more desiccated in cultivarsin which the time to abscission was longer, (c) fertilizationhad little effect on the time to abscission in most cultivars,whereas the absence of fertilization prevented petal abscissionin a number of the small-diameter cultivars where it was replacedby flower abscission, and (d) cutting and placement in waterat 20 °C did not affect the time to abscission.Copyright1995, 1999 Academic Press Abscission, fertilization, flowers, petals, Rosa hybrida L., rose, water stress, carbohydrate stress     

Flower power: its association with bee power and floral functional morphology in papilionate legumes

Background and Aims

A test was made of the hypothesis that papilionate legume flowers filter pollinators according to their ability to exert strength to open flowers to access rewards. In addition, interactions with pollen vectors were expected to explain the structural complexity of the architecture of these flowers since operative flower strength may be determined by a combination of morphological traits which form part of an intrafloral functional module.

Methods

Six papilionate species were studied: Collaea argentina, Desmodium uncinatum, Galactia latisiliqua, Lathyrus odoratus, Spartium junceum and Tipuana tipu. Measurements were made of the strength needed to open keels and the strength that pollinators were capable of exerting. Morphological traits of all petals were also measured to determine which of them could be either mutually correlated or correlated with operative strength and moment of strength and participated in a functional module.

Key Results

It was observed that pollinators were capable in all cases of exerting forces higher and often several times higher than that needed to access floral rewards, and no association could be detected between floral operative strength and strength exerted by the corresponding pollinators. On the other hand, strong and significant correlations were found among morphometric traits and, of these, with operative strength and moment. This was particularly evident among traits of the keel and the wings, presumably involved in the functioning of the floral moveable mechanism.

Conclusions

Though visitors are often many times stronger than the operative strength of the flowers they pollinate, exceptionally weak bees such as Apis mellifera cannot open the strongest flowers. On the other hand, strong correlations among certain petal morphometric traits (particularly between the keel and wings) give support to the idea that an intrafloral module is associated with the functioning of the mechanism of these legume flowers. In addition, the highly significant correlations found across petals support the view of functional phenotypic integration transcending the ontogenetic organization of flower structure.     

Why do so many petals have conical epidermal cells?

Background

The conical epidermal cells found on the petals of most Angiosperm species are so widespread that they have been used as markers of petal identity, but their function has only been analysed in recent years. This review brings together diverse data on the role of these cells in pollination biology.

Scope

The published effects of conical cells on petal colour, petal reflexing, scent production, petal wettability and pollinator grip on the flower surface are considered. Of these factors, pollinator grip has been shown to be of most significance in the well-studied Antirrhinum majus/bumble-bee system. Published data on the relationship between epidermal cell morphology and floral temperature were limited, so an analysis of the effects of cell shape on floral temperature in Antirrhinum is presented here. Statistically significant warming by conical cells was not detected, although insignificant trends towards faster warming at dawn were found, and it was also found that flat-celled flowers could be warmer on warm days. The warming observed is less significant than that achieved by varying pigment content. However, the possibility that the effect of conical cells on temperature might be biologically significant in certain specific instances such as marginal habitats or weather conditions cannot be ruled out.

Conclusions

Conical epidermal cells can influence a diverse set of petal properties. The fitness benefits they provide to plants are likely to vary with pollinator and habitat, and models are now required to understand how these different factors interact.     

Graft‐accelerated virus‐induced gene silencing facilitates functional genomics in rose flowers

Rose has emerged as a model ornamental plant for studies of flower development, senescence, and morphology, as well as the metabolism of floral fragrances and colors.Virus-induced gene silencing(VIGS) has long been used in functional genomics studies of rose by vacuum infiltration of cuttings or seedlings with an Agrobacterium suspension carrying TRV-derived vectors. However, VIGS in rose flowers remains a challenge because of its low efficiency and long time to establish silencing. Here we present a novel and rapid VIGS method that can be used to analyze gene function in rose,called ‘graft-accelerated VIGS', where axil ary sprouts are cut from the rose plant and vacuum infiltrated with Agrobacterium. The inoculated scions are then grafted back onto the plants to flower and silencing phenotypes can be observed within 5 weeks, post-infiltration. Using this new method, we successfully silenced expression of the RhDFR_1, RhA G, and RhNUDX_1 in rose flowers, and affected their color, petal number, as well as fragrance, respectively. This grafting method will facilitate high-throughput functional analysis of genes in rose flowers. Importantly, it may also be applied to other woody species that are not currently amenable to VIGS by conventional leaf or plantlet/seedling infiltration methods.     

Engineering of the rose flavonoid biosynthetic pathway successfully generated blue-hued flowers accumulating delphinidin

Flower color is mainly determined by anthocyanins. Rosa hybrida lacks violet to blue flower varieties due to the absence of delphinidin-based anthocyanins, usually the major constituents of violet and blue flowers, because roses do not possess flavonoid 3',5'-hydoxylase (F3'5'H), a key enzyme for delphinidin biosynthesis. Other factors such as the presence of co-pigments and the vacuolar pH also affect flower color. We analyzed the flavonoid composition of hundreds of rose cultivars and measured the pH of their petal juice in order to select hosts of genetic transformation that would be suitable for the exclusive accumulation of delphinidin and the resulting color change toward blue. Expression of the viola F3'5'H gene in some of the selected cultivars resulted in the accumulation of a high percentage of delphinidin (up to 95%) and a novel bluish flower color. For more exclusive and dominant accumulation of delphinidin irrespective of the hosts, we down-regulated the endogenous dihydroflavonol 4-reductase (DFR) gene and overexpressed the Irisxhollandica DFR gene in addition to the viola F3'5'H gene in a rose cultivar. The resultant roses exclusively accumulated delphinidin in the petals, and the flowers had blue hues not achieved by hybridization breeding. Moreover, the ability for exclusive accumulation of delphinidin was inherited by the next generations.     

Correlation between number and position of floral organs in Arabidopsis

Background and Aims

The study of variation in number, position and type of floral organs may serve as a key to understanding the mechanisms underlying their variation, and will make it possible to improve the analysis of gene function in model plant species by means of a more accurate characterization of mutant phenotypes. The present analysis was carried out in order to understand the correlation between number and position of floral organs in Arabidopsis thaliana.

Methods

An analysis of number and position of organs in flowers of wild type as well as in a series of mutations with floral organ position alterations was carried out, using light and electron microscopy. Variation common to different genotypes was analysed by means of individual diagrams, upon which generalized diagrams depicting variation in number and position of organs, were built by superimposition.

Key Results and Conclusions

It is shown that in the Arabidopsis flower a correlation exists between positions of petals and sepals, as well as between positions of stamens and carpels, whereas the position of carpels does not seem to depend on number and position of petals and stamens. This suggests that the position of organs in the basal (sepals) and apical (carpels) parts of the flower are determined before that in the intermediate zone. This assumption is consistent with the results of mathematical modelling and is supposed to be the consequence of stem-cell activity in the flower.     

Transient gene expression in rose petals via <Emphasis Type="Italic">Agrobacterium</Emphasis> infiltration

The study of gene function in roses is hampered by the low efficiency of transformation systems and the long time span needed for the generation of transgenic plants. For some functional analyses, the transient expression of genes would be an efficient alternative. Based on current protocols for the transient expression of genes via the infiltration of Agrobacterium into plant tissues, we developed a transient expression system for rose petals. We used β-glucuronidase (GUS) as a marker gene to optimize several parameters with effects on GUS expression. The efficiency of expression was found to be dependent on the rose genotype, flower age, position of petals within a flower, Agrobacterium strain and temperature of co-cultivation. The highest GUS expression was recorded in petals of the middle whirls of half-bloomed flowers from cultivars of ‘Pariser Charme’ and ‘Marvel’.     

Novel aspects of the flowers and floral pigmentation of two Cleome species (Cleomaceae), C. hassleriana and C. serrulata

Palely pigmented inflorescences of cultivated Cleome hassleriana Chodat (spider flower) have a unique two-toned appearance shown in the present study to be due to a loss of petal pigmentation within 24 h of anthesis, accompanied by an equally unique loss of petal mass. A similar loss occurs in deeply pigmented petals but is less evident to the eye because of the high initial content due to the presence, in the petal mesophyll, of globular anthocyanic vacuolar inclusions (AVIs). Inflorescences of the wild species, Cleome serrulata Pursh. (Rocky Mountain bee flower) are also two-toned because of the deeper pink colour of the unopened bud. No AVIs were seen. The pink colour of the bee flower petals is due to the same five acylated cyanidin glycosides as those previously isolated from mauve petals of spider flower. The structural pattern of the spider flower anthocyanins is shared with at least three genera of the Brassicaceae.     

Over-expression of the Gerbera hybrida At-SOC1-like1 gene Gh-SOC1 leads to floral organ identity deterioration

Background and Aims

The family of MADS box genes is involved in a number of processes besides controlling floral development. In addition to supplying homeotic functions defined by the ABC model, they influence flowering time and transformation of vegetative meristem into inflorescence meristem, and have functions in roots and leaves. Three Gerbera hybrida At-SOC1-like genes (Gh-SOC1–Gh-SOC3) were identified among gerbera expressed sequence tags.

Methods

Evolutionary relationships between SOC1-like genes from gerbera and other plants were studied by phylogenetic analysis. The function of the gerbera gene Gh-SOC1 in gerbera floral development was studied using expression analysis, protein–protein interaction assays and reverse genetics. Transgenic gerbera lines over-expressing or downregulated for Gh-SOC1 were obtained using Agrobacterium transformation and investigated for their floral phenotype.

Key Results

Phylogenetic analysis revealed that the closest paralogues of At-SOC1 are Gh-SOC2 and Gh-SOC3. Gh-SOC1 is a more distantly related paralogue, grouping together with a number of other At-SOC1 paralogues from arabidopsis and other plant species. Gh-SOC1 is inflorescence abundant and no expression was seen in vegetative parts of the plant. Ectopic expression of Gh-SOC1 did not promote flowering, but disturbed the development of floral organs. The epidermal cells of ray flower petals appeared shorter and their shape was altered. The colour of ray flower petals differed from that of the wild-type petals by being darker red on the adaxial side and greenish on the abaxial surface. Several protein–protein interactions with other gerbera MADS domain proteins were identified.

Conclusions

The At-SOC1 paralogue in gerbera shows a floral abundant expression pattern. A late petal expression might indicate a role in the final stages of flower development. Over-expression of Gh-SOC1 led to partial loss of floral identity, but did not affect flowering time. Lines where Gh-SOC1 was downregulated did not show a phenotype. Several gerbera MADS domain proteins interacted with Gh-SOC1.     

Sites of ethylene production in the pollinated and unpollinated senescing carnation (Dianthus caryophyllus) inflorescence

Production of endogenous ethylene from the styles, ovary and petals of pollinated and unpollinated flowers of Dianthus caryophyllus L. was measured. The rate of ethylene production of cut, unpollinated flowers aged in water at 18°C was low until the onset of petal wilting, when a rapid surge of ethylene occurred in all tissues. The flower ethylene production was evolved mostly from the styles and petals. The bases of petals from unpollinated, senescing flowers evolved ethylene faster and sometimes earlier than the upper parts. Treatment of cut flowers with propylene, an ethylene analogue, accelerated wilting of flower petals and promoted endogenous ethylene production in all flower tissues. Pollination of intact flowers also promoted endogenous ethylene production and caused accelerated petal wilting within 2–3 days from pollination. Although the data are consistent with the hypothesis that ethylene forms a link between pollination of the style and petal wilting, in the unpollinated flower the style and petals can evolve a surge of ethylene independently of each other, about the time when the petals irreversibly wilt. The results are discussed in relation to the role of ethylene in flower senescence.