Do feather-degrading bacteria actually degrade feather colour? No significant effects of plumage microbiome modifications on feather colouration in wild great tits

Parasites are known to exert selective pressures on host life history traits since the energy and nutrients needed to mount an immune response are no longer available to invest in other functions. Bird feathers harbour numerous microorganisms, some of which are able to degrade feather keratin (keratinolytic microorganisms) and affect feather integrity and colouration in vitro. Although named “feather-degrading” microorganisms, experimental evidence for their effects on feathers of free-living birds is still lacking. Here, we tested whether (i) keratinolytic microorganisms can degrade feathers in vivo and thus modify the colour of feathers during the nesting period and (ii) whether feather microorganisms have a long-term effect on the investment in colouration of newly moulted feathers. We designed treatments to either favour or inhibit bacterial growth, thus experimentally modifying plumage bacterial communities, in a wild breeding population of great tits (Parus major). Our analyses revealed no significant effects of the treatments on feather colours. Moreover, we found that differences in bacterial exposure during nesting did not significantly affect the colouration of newly moulted feathers. Our results suggest that significant feather degradation obtained during in vitro studies could have led to an overestimation of the potential of keratinolytic microorganisms to shape feather colouration in free-living birds.     

Fifty shades of white: how white feather brightness differs among species

White colouration is a common and important component of animal visual signalling and camouflage, but how and why it varies across species is poorly understood. White is produced by wavelength-independent and diffuse scattering of light by the internal structures of materials, where the degree of brightness is related to the amount of light scattered. Here, we investigated the morphological basis of brightness differences among unpigmented pennaceous regions of white body feathers across 61 bird species. Using phylogenetically controlled comparisons of reflectance and morphometric measurements, we show that brighter white feathers had larger and internally more complex barbs than duller white feathers. Higher brightness was also associated with more closely packed barbs and barbules, thicker and longer barbules, and rounder and less hollow barbs. Larger species tended to have brighter white feathers than smaller species because they had thicker and more complex barbs, but aquatic species were not significantly brighter than terrestrial species. As similar light scattering principals affect the brightness of chromatic signals, not just white colours, these findings help broaden our general understanding of the mechanisms that affect plumage brightness. Future studies should examine how feather layering on a bird’s body contributes to differences between brightness of white plumage patches within and across species.     

Temporal variation in size-assortative mating and male mate choice in a spider with amphisexual care

Males should be more selective when they have a high investment in reproduction, especially in species with biparental or paternal care. In this context, male mate choice can promote size-assortative mating (SAM) when (1) large males win intrasexual disputes, (2) large females are more fecund, and (3) males prefer larger females to smaller ones. In the spider Manogea porracea, males exhibit high reproductive investment by building their webs above those of females and exhibiting extended care of offspring in the absence of females. Under these circumstances, we expect the occurrence of SAM and male preference for large females. Herein, we performed observations and experiments in the field to evaluate the hypotheses that (1) M. porracea mates assortatively by size and (2) SAM is influenced by male mate choice. Furthermore, we measured variables that could affect mating patterns, the sex ratios, and densities of both sexes. Pairing in M. porracea was positively size-assortative in 2012, but not in 2013. Large males won most disputes for mates and preferred larger females, which produced more eggs. The inconsistency in detection of SAM was due to population dynamics, namely variations in sex ratio and population density across the breeding season. Furthermore, we found that the significance of male mate choice on sexual selection of body size in M. porracea strongly depends on the competition intensity for mating opportunities. The traditional sexual selection hypothesis of SAM needs to be reviewed and must include measures of competition intensity.     

Soil and preen waxes influence the expression of carotenoid-based plumage coloration

The signaling function of carotenoid-based plumage is mainly determined by the concentration of pigments in feathers. For this reason, most studies of the proximate control of coloration focus on processes during and preceding moult. In great tits Parus major, past research demonstrates that carotenoid-based plumage coloration honestly indicates male quality and, thus, may be a sexually selected signal. In this study, we investigate how dirt and preen oil influence the coloration of carotenoid-based feathers in the great tit. We collected six feathers from each individual bird; three feathers served as controls while the remaining three feathers were washed with a chloroform/methanol mixture to remove soil and preen waxes. We assessed plumage coloration using digital photography. This cleaning procedure slightly enhanced ornamentation; the experimentally cleaned feathers expressed hues shifted towards shorter wavelengths and expressed brighter overall coloration than control feathers. This is the first experimental study conducted on wild birds demonstrating that, in addition to pigment concentration, the presence of preen waxes and soils on feathers may contribute to variation in coloration.     

The effect of growth rate and ageing on colour variation of European pond turtles

Many chelonians have colourful dots, patches and stripes throughout their body that are made up, at least in part, of carotenoids. Therefore, turtles are very suitable models to study the evolution and functionality of carotenoid-based colouration. Recent studies suggested a close link between colouration and immune system in these taxa. However, more research is needed to understand the role of these colourful stripes and patches in turtle visual signalling. The purpose of this study was to explore the relationship between growth rate and colouration in European pond turtles. In particular, we wanted to answer the question of whether there is a trade-off between growth and colour expression. We also aimed to explore the effect of body size and age on colour variation. Turtles from a reintroduction-breeding program were recaptured, weighed and measured over an 8-year period to estimate their growth rates and age. We also measured with a spectrometer the reflectance of colour patches in two different body parts: shell and forelimb. We found that turtles with a faster growth rate had brighter limb stripes independently of their age. On the other hand, shell colouration was related to body size with larger turtles having brighter shell stripes and higher values of carotenoid chroma. Our results suggest that fast-growers may afford to express intense colourful limb stripes likely due to their higher intake of carotenoids that would modulate both growth and colour expression. However, shell colouration was related to body size probably due to ontogenetic differences in the diet, as juveniles are strictly carnivorous while adults are omnivorous. Alternatively, shell colouration might be involved in crypsis as the shell is visually exposed to predators.     

Carotenoids,immune response and the expression of sexual ornaments in male greenfinches (<Emphasis Type="Italic">Carduelis chloris</Emphasis>)

Allocation trade-offs of carotenoids between their use in the immune system and production of sexual ornaments have been suggested as a proximate mechanism maintaining honesty of sexual signals. To test this idea, we experimentally examined whether carotenoid availability in the diet was related to variation in antibody response to novel antigens in male greenfinches (Carduelis chloris aurantiiventris), a species with extensive carotenoid-dependent plumage colouration. We also measured the cost of mounting a humoral response in terms of circulating carotenoids. Finally, we examined the relationship between plumage colour, immune response and circulating carotenoids. We found that males with carotenoid-supplemented diets showed stronger antibody response than non-supplemented birds. We also found that activation of the immune system significantly reduced circulating carotenoids (24.9% lower in immune-challenged birds than in control birds). Finally, intensity (chroma) of ventral plumage colouration of males, a character directly related to concentration of total carotenoids in feathers, was negatively correlated with the immune response and circulating carotenoids in winter. These results support the idea that carotenoids are a limiting resource and that males trade ornamental colouration against immune response.     

Why do <Emphasis Type="Italic">Manduca sexta</Emphasis> feed from white flowers? Innate and learnt colour preferences in a hawkmoth

Flower colour is an important signal used by flowering plants to attract pollinators. Many anthophilous insects have an innate colour preference that is displayed during their first foraging bouts and which could help them locate their first nectar reward. Nevertheless, learning capabilities allow insects to switch their colour preferences with experience and thus, to track variation in floral nectar availability. Manduca sexta, a crepuscular hawkmoth widely studied as a model system for sensory physiology and behaviour, visits mostly white, night-blooming flowers lacking UV reflectance throughout its range in the Americas. Nevertheless, the spectral sensitivity of the feeding behaviour of naïve moths shows a narrow peak around 450 nm wavelengths, suggesting an innate preference for the colour blue. Under more natural conditions (i.e. broader wavelength reflectance) than in previous studies, we used dual choice experiments with blue- and white-coloured feeders to investigate the innate preference of naïve moths and trained different groups to each colour to evaluate their learning capabilities. We confirmed the innate preference of M. sexta for blue and found that these moths were able to switch colour preferences after training experience. These results unequivocally demonstrate that M. sexta moths innately prefer blue when presented against white flower models and offer novel experimental evidence supporting the hypothesis that learning capabilities could be involved in their foraging preferences, including their widely observed attraction to white flowers in nature.     

Assortative mating by carotenoid-based plumage colour: a quality indicator in American goldfinches,<Emphasis Type="Italic"> Carduelis tristis</Emphasis>

We investigated the mating patterns in 22 breeding pairs of wild American goldfinches (Carduelis tristis) with respect to their body size, condition, and carotenoid-based, yellow plumage colour. Using reflectance spectrometry, we objectively quantified plumage colours across the bird-visible wavelengths, revealing a unexpected UV peak in the reflectance spectrum from yellow feathers. We summarized our colour measurements using a principal components analysis to create a single variable, carotenoid PC1, that represents the intensity of this carotenoid-based yellow colour, a measure of phenotypic quality in this species. We found no evidence of assortative mating with respect to measures of body size or condition but there was positive assortative mating by carotenoid PC1, such that the yellow plumage colours of males and females were significantly correlated within pairs. We argue that the yellow carotenoid coloration of goldfinches may be important in mutual mate choice and, thus, that sexual selection in this species may act upon female ornamentation, as well as the more obvious plumage signals of males. Because assortative mating results in an increase in genetic variance, we suggest that this might be a mechanism that maintains variance in ornamental traits in spite of the variance-eroding effects of sexual selection.     

One trait, many signals: different information on male quality is enclosed within the same trait in a blenny fish

Colourful ornaments are traditionally evaluated as one trait. However, they could consist of several components, such as total size, colour intensity and extension, each possibly bearing its own message about one aspect of quality. Males of the blenny Salaria pavo exhibit a colourful head crest and solely care for eggs. During the breeding season, the head crest shows a yellow colouration, the intensity and relative extension of which are independent of crest size. Here, we show that: (1) carotenoids are responsible for the head crest yellow patch; (2) activating the immune system by injecting the bacterial antigen lipopolysaccharides affects both the intensity and extent of the yellow colouration; and (3) females assess males on the basis of colour patch expression. However, the response of the yellow patch to the immune challenge was dependent on head crest size. Indeed, males with a larger head crest reacted better to the simulated infection, sustaining a level of yellow patch close to pre-challenge size.     

The fractal dimension of a conspicuous ornament varies with mating status and shows assortative mating in wild red-legged partridges (<Emphasis Type="Italic">Alectoris rufa</Emphasis>)

Complex body designs, such as plumage ornaments in birds, can be described by fractal geometry. These complex patterns could have a role as visual signals during courtship and social interactions, but an empirical validation in the wild is currently lacking. Here, we investigated whether the fractal dimension (FD) of a complex plumage pattern displayed by red-legged partridges Alectoris rufa could function as a potential sexual signal. We captured wild birds early in the breeding season and tested if mated and unmated birds differed in the FD of their conspicuous melanin-based black bib. We also tested if the FD of the black bib was correlated within the pair, looking for evidence of assortative mating based on the expression of this trait. We simultaneously assessed similar effects in other ornamental traits (black bib size, white throat patch and black flank band surface, redness of the eye rings and bill). Mated birds showed higher black bib FD values than unmated ones. Mated males, but not females, also displayed a larger black bib. Moreover, the black bib FD (but not the trait size) and the white throat patch surface showed assortative mating. Finally, females with higher black bib FD showed smaller black flank band surface, suggesting a trade-off in the expression of the two melanin-pigmented plumage traits. This provides unique and novel indication for the shape complexity of a pigmented trait, here described by its fractal dimension, to be potentially under sexual selection in a wild animal.     

Gaps in the capacity of modern forage crops to adapt to the changing climate in northern Europe

The within-species diversity in response to weather and the gaps in the response diversity in the modern set of forage crop cultivars were determined using an approach that assessed the adaptive capacity under global climate change. The annual dry matter (DM) yields were recorded in multi-location MTT (Maa- ja elintarviketalouden tutkimuskeskus) Agrifood Research Official Variety Trials in Finland for modern forage crop cultivars from 2000 to 2012, as a response to agroclimatic variables critical to yield based on the year-round weather data. The effect and interaction of cultivars and agroclimatic variables were analysed using mixed model. The relatively low adaptive capacity of timothy (Phleum pratense L.) and meadow fescue (Festuca pratensis Huds.) indicates that diversification of the breeding material is warranted, particularly for resistance to high temperatures during primary growth and to high temperature sum 7 days after the first cut. All red clover cultivars (Trifolium pratense L.) suffered from both low and high accumulation of warm winter temperatures. Except for the red clover cultivars, cold stress during winter and lack of warm winter temperatures consistently reduced the yields of all species and cultivars. All tall fescue (Festuca arundinacea Schreb.) cultivars suffered from low precipitation during the fall hardening period. Although the set of festulolium (Festulolium pabulare) cultivars was also sensitive to low precipitation during the fall, festulolium was a good example of enhanced capacity to adapt to climate change with high response diversity because the cultivar germplasm base was diversified. Foreign origin in a cultivar pool was apparently not sufficient or necessary to ensure added value for a diversity of responses to climate change. Similar analyses to those used in this study, applied as practical tools for breeders, farmers and public actors, are important to secure the adaptive capacity of crops worldwide under global climate change.     

Optical properties of the uropygial gland secretion: no evidence for UV cosmetics in birds

Ultraviolet (UV) reflectance of the plumage is common in birds and plays an important role in sexual signalling. Recently, it has been proposed that birds are able to modify plumage UV reflectance by the application of uropygial gland secretion. Based on a survey of the optical properties of this secretion from 51 species belonging to 12 avian orders, we show that two main types of uropygial secretions exist, one predominantly found in passerines and one in non-passerines, both reducing relative UV reflectance of a white background (Teflon™ tape). We quantified how each type of secretion (exemplified by blue tit and mallard) affected feather UV reflectance. Both secretions reduced overall brightness and relative UV reflectance of white mallard feathers but hardly affected the reflectance of UV/blue blue tit crown feathers. According to models of avian colour vision, changes in reflectance due to application of the secretion were at or below the discrimination threshold of most birds. We conclude that the uropygial secretion is unlikely to play a major role in modifying plumage UV reflectance. However, the optical properties of the uropygial secretion may have been selected to interfere as little as possible with visual signaling through plumage reflectance. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Three-year lifecycle,large body,and very high threshold temperature in the cricket <Emphasis Type="Italic">Gryllus argenteus</Emphasis> for special adaptation to desiccation cycle in Malawi

In temperate climates, the initiation and termination of diapause synchronize the stress-tolerant stage with the stressful season and reproduction with the non-stressful season in many insects. Synchronization is often regulated by photoperiodism.Voltinism and the ultimate size of adults are also important determinants for their lifecycle, and different diapause stages and voltinism patterns are known in crickets.Here, we investigated the life history of the African cricket Gryllus argenteus from Malawi, which is a typical arid tropical highland. The climate is characterized by alternating arid and wet seasons, each of which lasts for half a year, and where the available heat mass is much less than lowlands at the same latitude. We first measured the nymphal duration at each rearing temperature and calculated the lower developmental threshold (t ₀) to be 20.19 °C based on Ikemoto and Takai (2000) and 19.38 °C based on a conventional line-fitting method. These values are very high relative to many other insects. The local temperature in winter does not fall below 15 °C, but this is much higher than the lethal limit. This suggested that critical stress in this locality was not coldness but low precipitation in winter. We estimated, based both on local temperature change and the Ikemoto and Takai’s t ₀, that G. argenteus required 3 years to complete its lifecycle unlike wet lowland species, where univoltinism or multi-voltinism are commonplace. Photoperiodism was observed in this species, but due to a lag between annual cycles in photoperiod, temperature, and humidity, photoperiodism alone cannot atune their lifecycle with local conditions.Synchronization in this species was achieved by three different adaptations: photoperiodism, high t ₀, and large body size, which give it a long lifecycle. Although the species cannot achieve a univoltine lifecycle because of its high t₀ value, it can escape from dry season by entering diapause at moderate temperatures, probably thereby achieving adaptive synchrony of lifecycle with both favorable and unfavorable seasons. A comparison between a conventional photothermogram and a newly formulated photohydrogram or photohygrogram demonstrates that even though sufficient heat is available, scarcity of water and thus scarcity of foliage should force the cricket to maintain diapause at intermediate temperature. The results suggested that high t ₀, large body size, and multi-ennial lifecycle mutually affect each other and formulate a unique adaptation under such an extreme environment.     

Climatic influences on the breeding biology of the agile frog (<Emphasis Type="Italic">Rana dalmatina</Emphasis>)

Severe population declines of amphibians have been shown to be attributed to climate change. Nevertheless, the various mechanisms through which climate can influence population dynamics of amphibians remain to be assessed, notably to disentangle the relative synergetic or antagonistic influences of temperature and precipitations on specific life history stages. We investigated the impact of rainfall and temperature on the egg-clutch abundance in a population of agile frog (Rana dalmatina) during 29 years (1987–2016) on 14 breeding sites located in Brenne Natural Park, France. Specifically, we examined the influence of environmental conditions occurring during five temporal windows of the year cycle corresponding to specific life history stages. Overall, our results suggest that the year-to-year fluctuations of egg-clutch abundances in Brenne Natural Park were partly dependent on local climatic conditions (rainfall and temperature). Climate seemed to influence breeding frogs during the autumn-winter period preceding reproduction. Spring and summer conditions did not influence reproduction. Additionally, we failed to detect effects of climatic conditions on newly metamorphosed individuals. Other factors such as density dependence and inter-specific interactions with introduced predators are likely to play a significant role in reproduction dynamics of the studied frog populations.     

Parental care compromises feeding in the pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>)

Providing parental care is potentially costly. Costs can arise through elevated energy expenditure or from an increased risk of mortality. A cost of parental care can also occur because a parent is compromised in their ability to forage. We used pumpkinseed Lepomis gibbosus, a fish with an alternative male mating strategy, to test whether parental males differed in their feeding in comparison with females and cuckolder males. To address this question, we examined the stomach contents of female, cuckolder male, and parental male pumpkinseed during the breeding season over an entire diel cycle. We showed that parental males had a lower total weight of food in their stomachs in comparison with females, while cuckolder males did not. Parental males also had a lower weight and number of chironomids in their stomachs. The temporal pattern of feeding of parental males diverged from that of females, and they had a lower probability of pupal chironomids in their stomachs, which implies spatial segregation in foraging. Parental males had a greater probability of conspecific eggs in their stomachs than females, while the probability of egg cannibalism did not differ between cuckolder males and females. Overall, these finding meet predictions in accordance with an assumption that parental care and territoriality can compromise feeding.     

<Emphasis Type="Italic">Fusarium</Emphasis> head blight incidence and mycotoxin accumulation in three durum wheat cultivars in relation to sowing date and density

Durum wheat (Triticum turgidum var. durum) is an important crop in Europe, particularly in the Mediterranean countries. Fusarium head blight (FHB) is considered as one of the most damaging diseases, resulting in yield and quality reduction as well as contamination of grain with mycotoxins. Three winter durum wheat cultivars originating from Austria, Slovakia, and Poland were analyzed during 2012–2014 seasons for FHB incidence and Fusarium mycotoxin accumulation in harvested grain. Moreover, the effects of sowing density and delayed sowing date were evaluated in the climatic conditions of Southern Poland. Low disease severity was observed in 2011/2012 in all durum wheat cultivars analyzed, and high FHB occurrence was recorded in 2012/2013 and 2013/2014 seasons. Fusarium graminearum was the most abundant pathogen, followed by Fusarium avenaceum. Through all three seasons, cultivar Komnata was the most susceptible to FHB and to mycotoxin accumulation, while cultivars Auradur and IS Pentadur showed less symptoms. High susceptibility of cv. Komnata was reflected by the number of Fusarium isolates and elevated mycotoxin (deoxynivalenol, zearalenone, and moniliformin) content in the grain of this cultivar across all three seasons. Nivalenol was identified in the samples of cv. Komnata only. Genotype-dependent differences in FHB susceptibility were observed for the plants sown at optimal date but not at delayed sowing date. It can be hypothesized that cultivars bred in Austria and Slovakia show less susceptibility towards FHB than the cultivar from Poland because of the environmental conditions allowing for more efficient selection of breeding materials.     

Breeding skylarks (Alauda arvensis) on organic and conventional arable farms in The Netherlands

The aim of this study was to analyse the effects of differences in cropping pattern between organic and conventional arable farms on the breeding activity of skylarks and to assess the effects of arable crop management on skylark nest survival. Skylark nest density was seven times higher on organic farms than on conventional farms (0.63 vs. 0.09 nest per 10 ha). Skylarks showed a strong preference for spring cereals, lucerne and grass leys, all of which were mainly or exclusively grown on organic farms. On organic farms nests were initiated during the entire breeding season, but on conventional farms no nesting activity was found during the peak of the season (early May to early June). On organic farms 27% of all nests was successful. Increasing the availability of suitable breeding habitat during the peak of the breeding season on conventional farms might provide one means of enhancing breeding skylark populations. On organic farms, crop management should focus on reducing nest loss due to farming operations.     

A comparative analysis of colour preferences in temperate and tropical social bees

The spontaneous occurrence of colour preferences without learning has been demonstrated in several insect species; however, the underlying mechanisms are still not understood. Here, we use a comparative approach to investigate spontaneous and learned colour preferences in foraging bees of two tropical and one temperate species. We hypothesised that tropical bees utilise different sets of plants and therefore might differ in their spontaneous colour preferences. We tested colour-naive bees and foragers from colonies that had been enclosed in large flight cages for a long time. Bees were shortly trained with triplets of neutral, UV-grey stimuli placed randomly at eight locations on a black training disk to induce foraging motivation. During unrewarded tests, the bees’ responses to eight colours were video-recorded. Bees explored all colours and displayed an overall preference for colours dominated by long or short wavelengths, rather than a single colour stimulus. Naive Apis cerana and Bombus terrestris showed similar choices. Both inspected long-wavelength stimuli more than short-wavelength stimuli, whilst responses of the tropical stingless bee Tetragonula iridipennis differed, suggesting that resource partitioning could be a determinant of spontaneous colour preferences. Reward on an unsaturated yellow colour shifted the bees’ preference curves as predicted, which is in line with previous findings that brief colour experience overrides the expression of spontaneous preferences. We conclude that rather than determining foraging behaviour in inflexible ways, spontaneous colour preferences vary depending on experimental settings and reflect potential biases in mechanisms of learning and decision-making in pollinating insects.     

Biological treatment of chicken feather waste for improved biogas production

A two-stage system was developed which combines the biological degradation of keratin-rich waste with the production of biogas. Chicken feather waste was treated biologically with a recombinant Bacillus megaterium strain showing keratinase activity prior to biogas production. Chopped, autoclaved chicken feathers (4%, W/V) were completely degraded, resulting in a yellowish fermentation broth with a level of 0.51 mg/mL soluble proteins after 8 days of cultivation of the recombinant strain. During the subsequent anaerobic batch digestion experiments, methane production of 0.35 Nm³/kg dry feathers (i.e., 0.4 Nm³/kg volatile solids of feathers), corresponding to 80% of the theoretical value on proteins, was achieved from the feather hydrolyzates, independently of the pre-hydrolysis time period of 1, 2 or 8 days. Cultivation with a native keratinase producing strain, Bacillus licheniformis resulted in only 0.25 mg/mL soluble proteins in the feather hydrolyzate, which then was digested achieving a maximum accumulated methane production of 0.31 Nm³/kg dry feathers. Feather hydrolyzates treated with the wild type B. megaterium produced 0.21 Nm³ CH₄/kg dry feathers as maximum yield.     

Testing the influence of gravity on flower symmetry in five <Emphasis Type="Italic">Saxifraga</Emphasis> species

Flower symmetry is considered a species-specific trait and is categorized in asymmetry, actinomorphic symmetry, bisymmetry and zygomorphic symmetry. Here we report on the intra-individual variation of flower symmetry in the genus Saxifraga and the influence of light, gravity and intrinsic factors on the development of flower symmetry. We tested five species—Saxifraga cuneifolia, Saxifraga imparilis, Saxifraga rotundifolia, Saxifraga stolonifera and Saxifraga umbrosa—concerning six flower parameters—angles between petals, petal length, petal pigmentation, angular position of carpels, movement of stamens and (only for S. imparilis and S. stolonifera) the length of the two lower elongated petals in regard to their position towards the stem. Specimens of all species were tested on a vertical clinostat as a gravity compensator, on a horizontal clinostat as a light incidence compensator and on a stationary control. The results show that the angle of incident light has no apparent impact on flower symmetry, whereas gravity affects the angular position of petals in S. cuneifolia and S. umbrosa and the petal colouration in S. rotundifolia. In S. cuneifolia and S. umbrosa, the absence of directional gravity resulted in the development of actinomorphic flowers, whereas the corresponding control flowers were zygomorphic. The development of flowers in S. rotundifolia was not altered by this treatment. The length of the two elongated petals in S. stolonifera and S. imparilis was not affected by gravity, but rather was determined by position of the flower within the inflorescence and resulted in asymmetrical flowers.