Hermine Alexandre’s research while affiliated with Université de Montréal and other places

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Publications (9)


The two parental species studied
The small maps at the bottom-right corner of the images indicate their geographic origin. Photo credits: S. Joly.
Schematic depiction of the phenotypic traits used in this study
For each multivariate trait, the whole range of corolla shape variation is represented by the landmarks (red dots) and semi-landmarks (black dots) of the corolla profile; the minimum and maximum shapes correspond to the most extreme values observed in the corresponding Principal Component Analysis (PCA) (see [63]). Under the univariate traits is an example of how measurements were performed from landmarks positions on a specific floral shape from landmarks (red dots).
Morphological variation (diagonal) for the traits in the F2 hybrid population and pairwise scatterplots showing their correlation (lower diagonal)
The yellow and red vertical lines on the histograms show the position of the parents, R. auriculatum and R. rupincola, respectively. Regression slopes are shown for traits comparisons with a significant Pearson correlation test (p < 0.05).
Comparative transcriptome sequence analysis of synonymous and non-synonymous nucleotide substitutions of the three Rhytidophyllum species
A) Plot of dN/dS ratio for each pairwise species comparisons. The line indicates the 1:1 slope and blue points represent open reading frames (ORFs) that have a dN/dS > 1. B) Venn diagram showing the ORFs that were found significant in the different pairwise comparisons in A.
Linkage map with 12 QTLs
The linkage groups (LG) were given the same names as in Alexandre et al. [63] and QTLs are represented by coloured bars to the right of their designated linkage groups, with letters on top referring to Table 2. The patterns in QTLs bars represent different significance thresholds indicated in the legend, and the horizontal bar represents the peak location (see Table 2). The 24 candidate genes are in red, including CYC and RAD from Alexandre et al. [63]. Positions of markers are in cM. Minor QTLs are those that were detected after including the primary QTL as a covariate in the model.

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Testing candidate genes linked to corolla shape variation of a pollinator shift in Rhytidophyllum (Gesneriaceae)
  • Article
  • Full-text available

July 2022

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83 Reads

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1 Citation

Valérie Poulin

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Delase Amesefe

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Floral adaptations to specific pollinators like corolla shape variation often result in reproductive isolation and thus speciation. But despite their ecological importance, the genetic bases of corolla shape transitions are still poorly understood, especially outside model species. Hence, our goal was to identify candidate genes potentially involved in corolla shape variation between two closely related species of the Rhytidophyllum genus (Gesneriaceae family) from the Antilles with contrasting pollination strategies. Rhytidophyllum rupincola has a tubular corolla and is strictly pollinated by hummingbirds, whereas R. auriculatum has more open flowers and is pollinated by hummingbirds, bats, and insects. We surveyed the literature and used a comparative transcriptome sequence analysis of synonymous and non-synonymous nucleotide substitutions to obtain a list of genes that could explain floral variation between R. auriculatum and R. rupincola. We then tested their association with corolla shape variation using QTL mapping in a F2 hybrid population. Out of 28 genes tested, three were found to be good candidates because of a strong association with corolla shape: RADIALIS, GLOBOSA, and JAGGED. Although the role of these genes in Rhytidophyllum corolla shape variation remains to be confirmed, these findings are a first step towards identifying the genes that have been under selection by pollinators and thus involved in reproductive isolation and speciation in this genus.

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Table 1 : Number of transitions between the different pollination strategies according to the stochastic mapping. The median values obtained from the character simulations over the posterior distribution of species tree are reported as well as 95% credible intervals. Ancestral states are in rows. 
Figure 3: Species phylogeny showing mean corolla shapes (after Procrustes analysis). Pollination strategies are shown with those that have been confirmed indicated by a black contour. Pie charts represent the joint probability of each state at nodes as estimated by stochastic mapping from only species with confirmed pollinators. Clade posterior probabilities are shown above branches. Outgroup taxa are not shown. 
Figure 4: Corolla shape morphospaces obtained from principal component analyses performed on (A) all species or on (B) species excluding hummingbird (both confirmed and inferred) and moth pollinated species. The large dots on the plot represent the species means, which are connected by a line to the floral shapes of the individuals belonging to the species (small dots). Thin-plate spline deformation grids show corolla shape variation along the principal components (plus or minus 2 standard deviation from the mean shape). Bellonia spinosa (bee pollinated) was not included in the morphometric analyses because it has a radial symmetry. 
Greater pollination generalization is not associated with reduced constraints on corolla shape in Antillean plants

December 2017

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323 Reads

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32 Citations

Evolution

Flowers show important structural variation as reproductive organs but the evolutionary forces underlying this diversity are still poorly understood. In animal-pollinated species, flower shape is strongly fashioned by selection imposed by pollinators, which is expected to vary according to guilds of effective pollinators. Using the Antillean subtribe Gesneriinae (Gesneriaceae), we tested the hypothesis that pollination specialists pollinated by one functional type of pollinator have maintained more similar corolla shapes through time due to more constant and stronger selection constraints compared to species with more generalist pollination strategies. Using geometric morphometrics and evolutionary models, we showed that the corolla of hummingbird specialists, bat specialists, and species with a mixed-pollination strategy (pollinated by hummingbirds and bats; thus a more generalist strategy) have distinct shapes and that these shapes have evolved under evolutionary constraints. However, we did not find support for greater disparity in corolla shape of more generalist species. This could be because the corolla shape of more generalist species in subtribe Gesneriinae, which has evolved multiple times, is finely adapted to be effectively pollinated by both bats and hummingbirds. These results suggest that ecological generalization is not necessarily associated with relaxed selection constraints.


Representation of realized abiotic niches of plants harbouring different pollination strategies. Axes e1 and e2 represent abiotic environmental variables. The three type of plants have the same fundamental abiotic niche (in orange) but their realized niches correspond to the overlap of their own fundamental niche and that of their pollinators (abiotic niche of pollinators in red for hummingbirds and green for bats). Consequently, if pollinator functional groups have different abiotic niches, plants relying on different pollinators should occupy different abiotic realized niches.
Phylogeny of Gesneria and Rhytidophyllum. Numbers over the branches represent posterior probabilities. To the right of the phylogeny, the first column corresponds to the geographical distribution of each species (orange: presence, grey: absence, LA: present in the Lesser Antilles). The second column represents pollination mode (question mark: unknown, green: bat specialist, purple: mixed strategy bat–hummingbird, pink: hummingbird specialist; asterisks indicate species for which the pollination mode has been inferred from morphological data; see the electronic supplementary material for details). The third column represent floral phenotypes (photos from John Clark and Simon Joly).
Bioclimatic niches of plants and pollinators. (a) Hummingbird specialists, (b) bat specialists, (c) generalists, (d) hummingbirds and (e) bats. (f) The projection of the pixels of the major islands and Lesser Antilles on the PCA. The grey squares in graphs (a–e) represent available bioclimatic condition over the Antilles (pixels of all the regions).
Niche identity (point) and range (line) of plant (hummingbird specialists, bat specialists and generalists) and pollinator (hummingbirds, bats) species over the first and second principal components. The mean and range of each major island and the Lesser Antilles are also shown (figure 3 for islands’ colour code).
Bioclimatic niches are conserved and unrelated to pollination syndromes in Antillean Gesneriaceae

November 2017

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147 Reads

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9 Citations

The study of the evolution of abiotic niches can be informative regarding the speciation drivers in a given group. Yet, two factors that could potentially affect niche evolution have seldom been addressed concomitantly, which are biotic interactions and geographical isolation. In this study, we used as a model group the Antillean plant genera Gesneria and Rhytidophyllum (Gesneriaceae) to evaluate the effect of pollinators and geographical isolation on the bioclimatic niche. These genera possess species characterized by interspecific geographical isolation in different islands and are pollinated by different pollinators. Some species are pollinated by hummingbirds, other by bats, while some are more generalists and are pollinated by pollinators from both functional groups. After describing the bioclimatic niches of plant species, we measured niche overlap for species pairs and we fitted Brownian motion and Ornstein-Uhlenbeck (OU) evolution models with multiple evolutionary regimes to test for an effect of pollination strategy or geographical isolation on bioclimatic niche evolution of these plants. The analysis of niche overlap between plant species, which could not be corrected for phylogenetic relationships, showed that it was significantly influenced by pollination mode and island distribution. By contrast, the best fitting evolutionary model on niche optima and tolerance was always an OU model with a unique selective regime, suggesting that neither pollination strategy nor island isolation had an important effect on bioclimatic niches at a macroevolutionary scale. Instead, we conclude that bioclimatic niches of Antillean Gesneriaceae evolved under phylogenetic conservatism and hypothesize that this macroevolutionary pattern could result from adaptation to temporally variable climates in the Antilles.



Increased pollination specialization did not increase corolla shape constraints in Antillean plants

October 2017

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23 Reads

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1 Citation

Flowers show important structural variation as reproductive organs but the evolutionary forces underlying this diversity are still poorly understood. In animal-pollinated species, flower shape is strongly fashioned by selection imposed by pollinators, which is expected to vary according to guilds of effective pollinators. Using the Antillean subtribe Gesneriinae (Gesneriaceae), we tested the hypothesis that pollination specialists pollinated by one functional type of pollinator have maintained more similar corolla shapes through time due to stronger selection constraints compared to species with more generalist pollination strategies. Using geometric morphometrics and evolutionary models, we showed that the corolla of hummingbird specialists, bat specialists, and species with a mixed-pollination strategy (pollinated by hummingbirds and bats; thus a more generalist strategy) have distinct shapes and that these shapes have evolved under evolutionary constraints. However, we did not find support for smaller disparity in corolla shape for hummingbird specialists compared to more generalist species. This could be because the corolla shape of more generalist species in subtribe Gesneriinae, which has evolved multiple times, is finely adapted to be effectively pollinated by both bats and hummingbirds. These results suggest that pollination generalization is not necessarily associated with relaxed selection constraints.


Table 1 Information about linkage groups. 
Figure 1 Measure of shape variation in the hybrid population and parents. (A) Flowers from both parents (top row), the self-pollinated F1 and samples from the F2 population; (B) position of landmarks on corolla pictures-red stars represent landmarks and small orange stars are semi-landmarks.
Figure 5 Distribution and correlation among traits in the hybrid population. Diagonal: traits distribution, the vertical lines correspond to the values of parent R. rupincola (red), parent R. auriculatum (green) and the self-pollinated F1 (purple). Lower left triangle, correlation among traits, if covariation is significant after Sidak correction, the regression line was plotted; Upper right triangle, regression coefficient, in bold if correlation is significant.
Figure 6 Linkage map and position of QTLs. QTLs positions are marked with 2-LOD confidence region, numbers right to the linkage groups represent markers position in cM. Unique names for QTLs correspond to the names given in Table 2.
Genetic architecture of pollination syndrome transition between hummingbird-specialist and generalist species in the genus Rhytidophyllum (Gesneriaceae)

June 2015

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286 Reads

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21 Citations

Adaptation to pollinators is a key factor of diversification in angiosperms. The Caribbean sister genera Rhytidophyllum and Gesneria present an important diversification of floral characters. Most of their species can be divided in two major pollination syndromes. Large-open flowers with pale colours and great amount of nectar represent the generalist syndrome, while the hummingbird-specialist syndrome corresponds to red tubular flowers with a less important nectar volume. Repeated convergent evolution toward the generalist syndrome in this group suggests that such transitions rely on few genes of moderate to large effect. To test this hypothesis, we built a linkage map and performed a QTL detection for divergent pollination syndrome traits by crossing one specimen of the generalist species Rhytidophyllum auriculatum with one specimen of the hummingbird pollinated R. rupincola. Using geometric morphometrics and univariate traits measurements, we found that floral shape among the second-generation hybrids is correlated with morphological variation observed between generalist and hummingbird-specialist species at the genus level. The QTL analysis showed that colour and nectar volume variation between syndromes involve each one major QTL while floral shape has a more complex genetic basis and rely on few genes of moderate effect. Finally, we did not detect any genetic linkage between the QTLs underlying those traits. This genetic independence of traits could have facilitated evolution toward optimal syndromes.


Figure 1. Measure of shape variation in the hybrid population and parents.
Table 2 .
Genetic architecture of pollination syndrome transition between hummingbird-specialist and generalist species in the genus Rhytidophyllum (Gesneriaceae)

May 2015

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16 Reads

Adaptation to pollinators is a key factor of diversification in angiosperms. The Caribbean sister genera Rhytidophyllum and Gesneria present an important diversification of floral characters. Most of their species can be divided in two major pollination syndromes. Large-open flowers with pale colours and great amount of nectar represent the generalist syndrome, while the hummingbird-specialist syndrome corresponds to red tubular flowers with a less important nectar volume. Repeated convergent evolution toward the generalist syndrome in this group suggests that such transitions rely on few genes of moderate to large effect. To test this hypothesis, we built a linkage map and performed a QTL detection for divergent pollination syndrome traits by crossing one specimen of the generalist species Rhytidophyllum auriculatum with one specimen of the hummingbird pollinated R. rupincola . Using geometric morphometrics and univariate traits measurements, we found that floral shape among the second-generation hybrids is correlated with morphological variation observed between generalist and hummingbird-specialist species at the genus level. The QTL analysis showed that colour and nectar volume variation between syndromes involve each one major QTL while floral shape has a more complex genetic basis and rely on few genes of moderate effect. Finally we did not detect any genetic linkage between the QTLs underlying those traits. This genetic independence of traits could have facilitated evolution toward optimal syndromes.


Figure 1. Measure of shape variation in the hybrid population and parents.
Table 2 .
Genetic architecture of pollination syndrome transition between hummingbird-specialist and generalist species in the genus Rhytidophyllum (Gesneriaceae)

May 2015

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22 Reads

Adaptation to pollinators is a key factor of diversification in angiosperms. The Caribbean sister genera Rhytidophyllum and Gesneria present an important diversification of floral characters. Most of their species can be divided in two major pollination syndromes. Large-open flowers with pale colours and great amount of nectar represent the generalist syndrome, while the hummingbird-specialist syndrome corresponds to red tubular flowers with a less important nectar volume. Repeated convergent evolution toward the generalist syndrome in this group suggests that such transitions rely on few genes of moderate to large effect. To test this hypothesis, we built a linkage map and performed a QTL detection for divergent pollination syndrome traits by crossing one specimen of the generalist species Rhytidophyllum auriculatum with one specimen of the hummingbird pollinated R. rupincola . Using geometric morphometrics and univariate traits measurements, we found that floral shape among the second-generation hybrids is correlated with morphological variation observed between generalist and hummingbird-specialist species at the genus level. The QTL analysis showed that colour and nectar volume variation between syndromes involve each one major QTL while floral shape has a more complex genetic basis and rely on few genes of moderate effect. Finally we did not detect any genetic linkage between the QTLs underlying those traits. This genetic independence of traits could have facilitated evolution toward optimal syndromes.


Citations (5)


... The expression pattern of CYC2-like genes has gradually evolved, and was widely expressed in the meristem of early-diverging Lamiales with a bilaterally symmetrical corolla, but limited in the meristem of core Lamiales and thus may be related to the origin of corolla bilateral symmetry [105,106]. The repeated loss of bilateral corolla symmetry is relatively frequent in Lamiaceae, which may be caused by different mechanisms and changes in floral symmetry-related genes, such as the loss of the CYC2 clade gene Ml-CYC2A in the genome and the contraction, expansion, or altered expression of Cc-CYC2A [107,108]. ...

Reference:

Advances in Research on the Regulation of Floral Development by CYC-like Genes
Testing candidate genes linked to corolla shape variation of a pollinator shift in Rhytidophyllum (Gesneriaceae)

... The most recent common plant ancestor was most likely a hummingbird specialist [34], and there have been several subsequent independent evolutions towards bat pollination, either keeping hummingbird pollination (in this case species have a mixed strategy) or losing it (species becoming bat specialists). Some reversals towards the ancestral hummingbird specialist mode were also detected but to a lesser extent [34,35]. The labile nature of pollination strategies gives the opportunity to study biological replicates of pollination transitions, making it easier to test for a link between pollination and abiotic niche evolution. ...

Increased pollination specialization did not increase corolla shape constraints in Antillean plants

... Ibañez et al. [52] investigated flower morphology variation in Jaborosa species, finding significant differences associated with pollination modes, particularly in sagittal view, suggesting adaptation to different pollinators. Similarly, Joly et al. [53] explored corolla shape variation in Gesneriinae, indicating distinct shapes among hummingbird specialists, bat specialists, and mixed-pollination species. Finally, Wolcott et al. [54] employed micro-CT and geometric morphometric analysis to study cacao's pollination biology, identifying the petal side door as a bottleneck for pollinator access and proposing new evidence for predicting unknown pollinators. ...

Greater pollination generalization is not associated with reduced constraints on corolla shape in Antillean plants

Evolution

... To summarize and visually inspect the available environmental conditions based on the 217 five filtered variables, we performed a principal component analysis (PCA) on the 455,929 land 218 pixels covering Hillieae's distribution (excluding the Caribbean and Atlantic forest) (Fig. 2c), 219 following Alexandre (Alexandre et al. 2017). We considered the first two axes of the PCA, 220 which explain most of the variation in the data. ...

Bioclimatic niches are conserved and unrelated to pollination syndromes in Antillean Gesneriaceae

... While this is true for the dimensions of the whole corolla of D. purpurea, our results suggest that both sections of the corolla behave as independent modules with different autonomy to respond to selection, with the proximal section (not directly involved in pollen transfer) being more evolutionarily flexible than the rest of the corolla. The genetic architecture of these traits in D. purpurea is unknown, but QTL studies in several lineages with shifts to hummingbird pollination show multiple loci of small or medium effect underlying the variation in corolla morphology (Stuurman et al., 2004;Wessinger et al., 2014;Alexandre et al., 2015;Kostyun et al., 2019), suggesting potential for complex responses. Regardless of the genetic basis, this decoupling between sections of the corolla might be derived from them being shaped by differential patterns of natural selection (to be described later). ...

Genetic architecture of pollination syndrome transition between hummingbird-specialist and generalist species in the genus Rhytidophyllum (Gesneriaceae)