of information on mating styles and reproductive fitness of theOpisthopappus , we cannot determine the likely causes for the
heterozygote excess and further research is needed.
4.2 Landscape factors determine gene exhchange pattern of
Opisthopappus
species
Maintaining an adequate migration rate is usually beneficial because the
populations of species experiencing limited gene flow would be
susceptible to loss of genetic diversity or genetic variation due to
inbreeding and drift (Bansept et al., 2021; Kottler et al., 2021). Lower
genetic diversity or genetic variation may make species difficult to
adapt to changing environmental conditions, leading to local extinctions
(Matzrafi et al., 2021; Wellband et al., 2021).
The gene exchange degree within Opisthopappus species present a
pattern mainly from north to south. The higher altitude of the northern
in Taihang mountains may contribute to some extent to the genes spread
of the Opisthopappus species. Meanwhile, the pollens and seeds ofOpisthopappus are small and light, and easily spread with wind.
Each year during the breeding period of the Opisthopappus , the
wind by Siberia weather in the Taihang Mountains could be an important
factor that facilitate the gene transmission from north to south (Wadl
et al., 2022).
Although there is gene exchange between two species and four groups, the
gene exchange was extremely limited (Fig. 4, 5 and Table S1). This
indicated the existence of a gene exchange barrier between O.
longilobus and O. taihangensis or among groups (Fig. 6).
Whilist, no obvious topographic obstacles were observed at the barrier
between Opisthopappus species. Thus, it suggested that the
factors hindering gene migration among populations might not be simple
topographic landscape factors but other factors of Taihang Mountians
landscape (Lu et al., 2021; Zhang et al., 2021). Taihang Mountains have
many deep gullies and diverse landforms, which could result in diverse
micro-landscape in different regions. This landscape heterogeneity could
produce heterogeneous habitats for different populations of species and
bring the spatial isolation for gene exchange.On the other hand, gene
exchange/flow is often considered a source of maladaptation because it
can limit genetic and phenotypic differentiation and reduce mean fitness
in a population (Youngblut et al., 2015). Limited gene exchange/flow
occurred in O. longilobus and O. taihangensis might be
benifical, even when populations are geographically disparate, because
gene flow can allow for the spread of environment-specific beneficial
alleles that arise in a single population that could bring the
differentiation and adaption (Ellstrand, 2014; Morente‐López et al.,
2021).
Meanwhile, the gene exchange barrier was located at the juncture of the
Hebei, Shanxi, and Henan provinces. The areas of this barrier were
suspected to be closely related to human activities. As the flowers ofOpisthopappus are beautiful and have good medicinal value, human
picking and planting activities may significantly influence migration
rates (Blanco et al., 2021; Crispo et al., 2011; Liao et al., 2021).
Among the landscape factors of Taihang Mountains, eight factors were the
key ones, mainly including six climatic factors and two human activity
factors. These factors might be heterogeneous in different regions and
form a climatic isolation and/or a corridor by human activites for the
populations of species, and which indicating that these factors are
important for the development and growth of O. longilobus andO. taihangensis (Ma et al., 2018; Shih et al., 2018).
For the two species, their flowering begin from June to September. And
their seeds develop during September and November. During the
development of breeding organs, they need accumulated temperature and
precipitation, thus four climatic factors, average precipitation in
August, average precipitation in October, average precipitation in
November, and solar radiation in August, would play an important role.
Except for the above factors, built-up land and rain-fed cultivated land
factors also significantly impacted the Opisthopappus (Table 3).
These two human activities are destructive to the environment that has a
long-term occupation and large-scale deep-level transformation on the
natural landscape especially physical and chemical properties of the
land, which would result in the changes of land workability and soil PH.
Based on the results of Samßada and LFMM, average precipitation in
November might have the greatest influence on the Opisthopappus(Shih et al., 2018). In the reciprocal transplant, it was found that too
much or too little precipitation would lead to seed growth with
cavitation. Therefore, it is speculated that the precipitation in
November may be regarded as a ecological indicator for theOpisthopappus two species (An et al., 2020; Chen et al., 2021;
Wang et al., 2010).
Meanwhile, among the pathways mainly enriched by the identified selected
SNPs, MAPK signaling and plant hormone signal transduction pathways were
significant ones. The two pathways are directly or indirectly related to
resistance of plant. In the growth and development of Arabidopsis
thaliana , it has been shown that MAPK signaling pathway was related to
environmental response (Fengbo, 2019). Plant hormone signal transduction
pathway has been shown to be related to the plant height through
affecting the cell elongation (Haruta and Sussman, 2017). Plant height
can affect the ability of plants to receive sunlight, which might be one
reason why solar radiation in August is an important factor forOpisthopappus species (Wang et al., 2021a). Thus, it could be
hypothesized that precipitation, solar light and soil PH might paly a
signal role that activate or inhibit the relative gene express, thereby,
bring the adaption of two species to the different landscapt in the
Taihang Mountains.
Conclusion
Landscape heterogeneity often plays a vital role in the ability of
organisms to disperse or not to disperse. Habitat fragmentation, both
natural and artificial, adversely affects the connectivity and
persistence of populations. Various landscape features of the Taihang
Mountains could make the different environmental conditions and
fragmentized habitat of each population or species ofOpisthopappus . With the changes of climate, and human activities,
landscape heterogeneity lead to the barrier among Opisthopappusspecies and hinder the gene exchange among the population and promote
the differentiation and adaptation among populations ever between
species.
CRediT authorship contribution
statement
Hao Zhang: Methodology, Writing – review & editing. Hang Ye:
Methodology, Editing. Li Liu: Methodology. En Zang: Analysis,
Investigation. Qiyang Qie: Sorting data, Investigation. Shan He:
Investigation. Weili Hao: Analysis, Investigation. Yafei Lan:
Investigation. Zhixia Liu: Investigation. Genlou Sun: Supervision,
Review & editing. Yiling Wang: Conceptualization, Funding acquisition,
Validation, Writing – review & editing.
Declaration of Competing
Interest
The authors declare that they do not have any competing financial
interests or personal relationships that could have appeared to
influence the work reported in this paper.
Acknowledgments
This work was supported by the National Natural Science Foundation of
China (31970358); Research Project of Shanxi Scholarship Council
(2020-090); Natural Science Foundation of Shanxi Province of China
(20210302124501); and Graduate Education nnovation Project of Shanxi
(2020SY323).
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