This thesis demonstrated the phylogeny, evolutionary history, genetic regulation of actinomorphic flower development and GCYC evolution in association with floral symmetry transition of
Conandron. The results from multi loci phylogeny (Fig. 1-3, Chapter 1) not only provide first evidence for the genetic distinctiveness of var. ramondioides and var. taiwaensis, but also support treatment of var. ryukuensis by Masamune (1939). Moreover, plants distributed in Southeast China also formed a geographical corresponding clade, suggesting a potential variety in Southeast China.
Our preliminary measurement of lobe-length to tube-length ratio from three sampled Southeast China Conandron flowers may support the treatment of plants from Southeast China (Fig. 5-1). By plotting the lobe-length to tube-length ratio of samples from Taiwan (collecting by me and Hong-wen Ma), Japan (measuring from specimen in TAI herbarium) and Southeast China (collecting by me and Dr. Chun-Neng Wang), three groups were identified. First, measurement of samples from Taiwan distributed relatively close to each other, forming one group. Similar pattern can also be indicated from measurement of Japan samples. Measurement of lobe-length to tube-length ratio of Taiwan and Japan samples is consistent with findings from Kokubugata and Peng (2004). However, measurement from Southeast China samples against the assumption proposed by Kokubugata and Peng (2004). Due to lack of samples from Southeast China, they can only judge from description of plants in Southeast China in China flora (Wang et al., 1998) and assigned plants in Southeast China into var. taiwanensis. However, based on measurement of Southeast China samples in this thesis, morphological trait obtaining form plants in Southeast China is close to that obtaining from Japan specimen instead of Taiwan samples. Discordance between my measurement and that proposed by Kokubugata and Peng (2004) suggests that collecting and measuring speciemen from Southeast China is necessary to resolve taxonomic treatment of Conandron distributed in Southeast China.
Phylogeographical studies of plants in East Asia tend to correlate lineage divergence process with geographical events (e.g. glacial and inter-glacial period). However, it is not the case in Conandron.
Instead, the reproductive characters of Conandron (e.g. lack of long distance dispersal structure on
seeds, limited pollen dispersal) play an important role in shaping lineage divergence process of this species (Chapter 1).
Applying molecular markers to infer is the current trend in studying evolutionary history of target species. However, evolutionary history of molecular markers may vary from each other. How to deal with discrepancy among molecular markers is an important issue before further analysis. In this thesis, discrepancy among molecular markers were identified between cpDNA and nuclear markers and among nuclear markers. Instead of ignoring discrepancy among molecular markers, methods were applied to clarify possible mechanisms in affecting evolutionary history of these genes in this thesis. For example, paml analysis was used to evaluate whether GCYC1C marker isolated from Conandron evolved neutrally or not. Identify neutrality of target markers is a key step before further analysis. That is because algorithms implemented in software or packages assume genetic
information obtained from molecular marker evolve neutrally. GCYC1, as a floral symmetry determining gene, could evolve under selection. Therefore, examining neutrality of GCYC1 is necessary. Examining results indicated GCYC1 in Conandron evolving non-neutrally in this thesis.
The non-neutral evolving results suggests using genetic information from GCYC1 may blur
evolutionary history of Conandron. For another nuclear marker, ITS, due to availability of universal primer, easy amplification and orthologue evolved concertedly, is broadly used in plant studies. The discrepancy between ITS and other nuclear marker makes us doubting. To examine whether ITS evolved concertedly, cloning methods and proof reading taq were used. Results indicated ITS isolated from Conandron evolved non-concertedly. Due to non-conceted evolution of ITS, ITS dataset was ignored in following analysis. As for discrepancy between cpDNA and nuclear markers (GCYC1, AGT intron 1, LFY intron 1 and GroES intron 1), non-neutral force may act on Southeast China i cpDNA, leading to non-monophyletic lineages of Southeast China cpDNA haplotypes in cpDNA haplotype network. By exploring possible mechanisms behind discrepancy among markers allows us to keep those neutral evolving markers and reconstruct a reliable phylogeny of Conandron.
This thesis demonstrated importance of identifying possible mechanism among molecular markers.
Filtering suitable markers for phylogeographical study is especially important in this era. That is because obtaining massive genetic data from interested species through Next generation sequencing is fast and convenient nowadays.
Establishment of actinomorphy in Conandron through loss of CYC expression in flower tissue suggesting the known CYC dependent regulatory pathway is flexible in association with floral symmetry transition (Chapter 2). This is been verified in other two Old World distributed
actinomorphic flower species, including Bournea (Zhou et al., 2008) and Tengia (Pang et al., 2010).
However, it is still unknown in New World distributed species, such as Niphaea oblonga. N. oblonga demonstrated entirely different floral trait from Conandron, Bournea and Tengia. In N. oblonga, the petal whorl exhibits actinomorphy at anthesis, but stamen whorl displays zygomorphy at anthesis.
That is because there is one aborted stamen located in dorsal region of flower, leading to
zygomorphy in stamen whorl of N. oblonga. The stamen-petal whorl exhibit difference symmetry type of N. oblonga is not seen in Conandron, Bournea and Tengia. By studying floral development of N. oblonga at molecular regulatory level, it is possible to find a novel modification of known CYC dependent regulatory pathway in association with actinomorphic flower establishment in N. oblonga.
In Chapter 3, relaxation of GCYC duplicated genes from genetic constrain is the possible
evolutionary force in maintaining multiple GCYC genes in Trichosporeae after duplication events occurred. However, the selection acting on the other two clades, Old World GCYC2 and New World GCYC1 clade, remains unknown. Between Old World GCYC2 and New World GCYC1, the New World GCYC1 clade is especially worth noting. There are four reasons. The first one is that ancestral symmetry form of New World species is zygomorphy (Chapter 3). The second reason is that
zygomorphy is dominant in New World species. The third reason is that the New World GCYC1 expressed in dorsal region of flower of Sinningia speciosa (Hsu et al., 2017). The forth reason is
GCYC1 isolated from New World distributed actinomorphic flower species containing putative full function GCYC1 gene (Smith et al., 2004). Judging from these four reasons, one question appears.
When reversal to actinomorphy occurred in a zygomorphic dominant clade, like in New World Gesneriaceae, what is the selection force behind? To answer this question, adding expression pattern of actinomorphic flower from New World species and comparing selection force between those actinomorphic flower lineages and zygomorphic flower lineages may help.
Figure
Fig. 5-1 Corolla lobe-length and tube-length plot of investigated Conandron individuals.
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8
0 0.2 0.4 0.6 0.8 1 1.2 1.4
averaged Tube length (cm)
averaged Lobe length (cm)
Taiwan Japan China
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