Polyploidy,or whole-genome duplication,is an important evolutionary process that has shaped the genomes and traits of many plants,including numerous important crops.The Brassica genus,which includes diverse vegetables...Polyploidy,or whole-genome duplication,is an important evolutionary process that has shaped the genomes and traits of many plants,including numerous important crops.The Brassica genus,which includes diverse vegetables and oilseeds,is a key model system for studying how polyploidy affects plant diversification and domestication.This review summarizes the current understanding of how multiple rounds of ancient and more recent polyploidization events laid the foundation for the wide diversity seen in Brassica.We discuss the key outcomes through which polyploidy facilitates the accumulation of genetic variation,including genomic buffering that enables mutation retention.Furthermore,we explore the significant roles of interspecies and interploidy introgression in introducing external genetic novelty.We highlight homoeologous exchange(HE)as a critical mechanism unique to allopolyploids,driving substantial genomic rearrangements including presence-absence variations and gene dosage alterations that directly contribute to significant phenotypic innovation and adaptation in Brassica.Together,these polyploidy-associated processes have led to the extensive range of genomic variations that shaped great morphological diversification in the domestication of Brassica.By integrating insights from genomics,genetics,and evolutionary biology,this review shows how polyploidy has been central to Brassica's success and agricultural value.We also suggest future research areas to better understand polyploid evolution and improve crop breeding.展开更多
[Objective] The aim was to study the feasibility of using colchicine to induce polyploidy from the buds of Shandong Salvia miltiorrhiza.[Method] The buds of Shandong Salvia miltiorrhiza were induced by colchicine.[Res...[Objective] The aim was to study the feasibility of using colchicine to induce polyploidy from the buds of Shandong Salvia miltiorrhiza.[Method] The buds of Shandong Salvia miltiorrhiza were induced by colchicine.[Result] When the concentration of colchicine supplemented in culture medium was 12 mg/L,the induction rate of polyploidy was the highest（36.6%）,but its survival rate was only 16.7%.When the concentration of colchicine was 8 mg/L,the induction rate of polyploidy was 15.9%,and its survival rate was up to 40.0%.Polyploidy plant of Shandong Salvia miltiorrhiza was strong and had thick,large and dark leaves,and long stomata on the lower epidermis of leaves had small density.[Conclusion] Colchicine could induce buds effectively,and the polyploidy breeding material of Salvia miltiorrhiza was obtained,which laid a foundation for shortening breeding process.展开更多
Dasiphora fruticosa comprises male,female and hermaphrodite plants,which are distributed sympatrically in some populations on the Qinghai-Tibet Plateau.To explore what governs the coexistence of these three sexual phe...Dasiphora fruticosa comprises male,female and hermaphrodite plants,which are distributed sympatrically in some populations on the Qinghai-Tibet Plateau.To explore what governs the coexistence of these three sexual phenotypes,we investigated the DNA contents,pollen and ovule production,pollen deposition,and performed hand-pollination in both hermaphroditic and dioecious individuals of D.fruticosa.Flow cytometry confirmed that the DNA content of males and females were almost twice as much as that of the hermaphrodites.Male and female flowers produced more pollen grains and ovules than hermaphroditic flowers.Hand-pollinated treatments showed that unisexual flowers were sterile in one sexual function and bisexual flowers were fertile for both functions,but no sterile seeds were produced between unisexual and bisexual flowers.Our findings imply that polyploidy is related to gender dimorphism,and both are likely to play a strong role in the coexistence of two cryptic biological species of D.fruticosa(low ploidy hermaphroditic species and high ploidy dioecious species)in the Qinghai-Tibet Plateau.展开更多
Possibility of polyploidy induction by colchicine in “Balady” mandarins was investigated in vitro. Seeds were immersed in different concentrations of colchicine solutions (0.01%, 0.05%, 0.1% and 0.2%) for different ...Possibility of polyploidy induction by colchicine in “Balady” mandarins was investigated in vitro. Seeds were immersed in different concentrations of colchicine solutions (0.01%, 0.05%, 0.1% and 0.2%) for different durations (12, 24 and 48 hr), then cultured in vitro on MS medium at half strength. Seed survival percentage decreased by increasing colchicine concentration and duration of treatment showing the lowest percentage at 0.2% for 48 hr. The highest DNA content was recorded at 0.2% for 24 hr. Stomata No. per unit area was decreased by colchicine treatments;moreover stomata length and width were studied. The results indicated that colchicine treatment at 0.1% for 48 hr had the highest tetraploid induction efficiency percentage.展开更多
Tea （Camellia sinensis L.） represents different ploidy levels. In the present paper, we reviewed the recent data on the diploid, aneuploid and polyploid formed their origin and chemically induced polyploidy and pred...Tea （Camellia sinensis L.） represents different ploidy levels. In the present paper, we reviewed the recent data on the diploid, aneuploid and polyploid formed their origin and chemically induced polyploidy and predicted it role in teagenetic improvement for better yield. Different polidy levels had different effects on tea physiology. Tetraploid and triploid had more vigour and hardness due to increased size of cells, while triploid could have even more vigours due to increased size of cells and sterilities. Chemically induced polyploidy had been found an important tool for improving plant physiology and production, therefore, induced polyploids should be produced to overcome the problem of low yield and limited rainfall in tea growing areas.展开更多
An ancient genome duplication (PPP1) that predates divergence of the cereals has recently been recognized. We report here another potentially older large-scale duplication (PPP2) event that predates monocot-dicot dive...An ancient genome duplication (PPP1) that predates divergence of the cereals has recently been recognized. We report here another potentially older large-scale duplication (PPP2) event that predates monocot-dicot divergence in the genome of rice (Oryza sativa L.), as inferred from the age distribution of pairs of duplicate genes based on recent genome data for rice. Our results suggest that paleopolyploidy was widespread and played an important role in the evolution of rice.展开更多
This study was planned to assess genetic variation in Jatropha curcas as affected by colchichine treatments. During the experiment, seeds of Jatropha curcas were collected from mature and healthy Jatropha plants and w...This study was planned to assess genetic variation in Jatropha curcas as affected by colchichine treatments. During the experiment, seeds of Jatropha curcas were collected from mature and healthy Jatropha plants and were soaked in different concentrations of colchicines. The second treatment was that of time of soaking (seeds were soaked in different concentrations for different time durations). The results clearly revealed that by increasing the concentration of colchicine treatment, parameters like stomatal density, and size of the guard cells were reduced.展开更多
Megakaryocytes and hepatocytes are unique cells in mammals that undergo polyploidization through endomitosis in terminal differentiation.Many polyploidization regulators and underlying mechanisms have been reported,mo...Megakaryocytes and hepatocytes are unique cells in mammals that undergo polyploidization through endomitosis in terminal differentiation.Many polyploidization regulators and underlying mechanisms have been reported,most of which are tightly coupled with development,organogenesis,and cell differentiation.However,the nature of endomitosis,which involves successful entry into and exit from mitosis without complete cytokinesis,has not yet been fully elucidated.We highlight that endomitosis is a new cell fate in the cell cycle,and tetraploidy is a critical stage at the bifurcation of cell fate decision.This review summarizes the recent research progress in this area and provides novel insights into how cells manipulate mitosis toward endomitosis.Endomitotic cells can evade the tetraploidy restrictions and proceed to multiple rounds of the cell cycle.This knowledge not only deepens our understanding of endomitosis as a fundamental biological process but also offers new perspectives on the physiological and pathophysiological implications of polyploidization.展开更多
Gerbera,a popular commercial cut flower with vibrant and striking colors has gained immense popularity in the floriculture industry.They are widely cultivated in various regions,making them available throughout the ye...Gerbera,a popular commercial cut flower with vibrant and striking colors has gained immense popularity in the floriculture industry.They are widely cultivated in various regions,making them available throughout the year.As a better alternative to conventional propagation methods(via seeds and rhizomes),plant tissue culture serves as way to avail large-scale,uniform,disease-free plantlets for commercial cultivation as well as to develop novel genotypes.In addition,it ensures production of healthy plantlets throughout the year in limited space.Based on the plant tissue culture techniques,the in vitro polyploidization,mutagenesis,and genetic transformation pave a path for creation of variation and eventually enhancing the ornamental traits to address the consumers’preferences and also facilitates in developing stress tolerant lines thereby minimizing the losses during cultivation,maintaining the quality of the flowers.This comprehensive review article presents an overview of the recent advancements on genetic improvement of gerbera via various cutting-edge plant tissue culture-based tools and techniques that contribute in enhancing the quality and efficiency of gerbera cultivation,meeting the demands of the floriculture industry while addressing the challenges of changing environment and resource limitations.展开更多
Dear Editor,Transcriptomic data have been widely employed to infer whole genome duplications (WGDs, or paleopolyploidy)(Jiao et al., 2011;Barker et al., 2016;Huang et al., 2016), through the use of large numbers of re...Dear Editor,Transcriptomic data have been widely employed to infer whole genome duplications (WGDs, or paleopolyploidy)(Jiao et al., 2011;Barker et al., 2016;Huang et al., 2016), through the use of large numbers of recognized paralogs with associated gene tree and Ks distributions, with the ability to date such polyploidy events.展开更多
Polyploidy refers to two or more complete sets of chromosomes that are combined within a nucleus,and includes both autopolyploidy and allopolyploidy(Otto 2007).Polyploidy is a recurring theme in the evolutionary histo...Polyploidy refers to two or more complete sets of chromosomes that are combined within a nucleus,and includes both autopolyploidy and allopolyploidy(Otto 2007).Polyploidy is a recurring theme in the evolutionary history of plants and it has been considered an important mechanism for speciation and also for adaptability to changing environments(Shimizu-Inatsugi et al.2017).In nature,about one-third of flowering plants are polyploids,and a great majority of them are allopolyploids.There are many cultivated allopolyploid species,such as cotton,Brassica and potato,the possible evolutionary history of which can be inferred with high certainty.展开更多
Polyploid cells,which contain more than one set of chromosome pairs,are very common in nature.Polyploidy can provide cells with several potential benefits over their diploid counterparts,including an increase in cell ...Polyploid cells,which contain more than one set of chromosome pairs,are very common in nature.Polyploidy can provide cells with several potential benefits over their diploid counterparts,including an increase in cell size,contributing to organ growth and tissue homeostasis,and improving cellular robustness via increased tolerance to genomic stress and apoptotic signals.Here,we focus on why polyploidy in the cell occurs and which stress responses and molecular signals trigger cells to become polyploid.Moreover,we discuss its crucial roles in cell growth and tissue regeneration in the heart,liver,and other tissues.展开更多
Cottons are the most important fiber crops in the world.The cotton genus Gossypium has 52 species,including seven allotetraploid species and 45 diploids.Four species were domesticated and remain as crops under cultiva...Cottons are the most important fiber crops in the world.The cotton genus Gossypium has 52 species,including seven allotetraploid species and 45 diploids.Four species were domesticated and remain as crops under cultivation today:the New World allopolyploid species G.hirsutum and G.barbadense(2n=52),and the Old World diploid species G.arboreum and G.herbaceum(2n=26).The primary cultivated species is Upland cotton(G.hirsutum L.),which accounts for more than 90%of global cotton fiber production.展开更多
Polyploidy is a major factor in the evolution of plants,yet we know little about the origin and evolution of polyploidy in intertidal species.This study aimed to identify the evolutionary transitions in three truemang...Polyploidy is a major factor in the evolution of plants,yet we know little about the origin and evolution of polyploidy in intertidal species.This study aimed to identify the evolutionary transitions in three truemangrove species of the genus Acanthus distributed in the Indo-West Pacific region.For this purpose,we took an integrative approach that combined data on morphology,cytology,climatic niche,phylogeny,and biogeography of 493 samples from 42 geographic sites.Our results show that the Acanthus ilicifolius lineage distributed east of the Thai-Malay Peninsula possesses a tetraploid karyotype,which is morphologically distinct from that of the lineage on the west side.The haplotype networks and phylogenetic trees for the chloroplast genome and eight nuclear genes reveal that the tetraploid species has two sub-genomes,one each from A.ilicifolius and A.ebracteatus,the paternal and maternal parents,respectively.Population structure analysis also supports the hybrid speciation history of the new tetraploid species.The two sub-genomes of the tetraploid species diverged from their diploid progenitors during the Pleistocene.Environmental niche models revealed that the tetraploid species not only occupied the near-entire niche space of the diploids,but also expanded into novel environments.Our findings suggest that A.ilicifolius species distributed on the east side of the Thai-Malay Peninsula should be regarded as a new species,A.tetraploideus,which originated from hybridization between A.ilicifolius and A.ebracteatus,followed by chromosome doubling.This is the first report of a true-mangrove allopolyploid species that can reproduce sexually and clonally reproduction,which explains the long-term adaptive potential of the species.展开更多
基金supported by the National Key Research and Development Program of China(Grant No.2021YFF1000104)the Agricultural Science and Technology Innovation Program(ASTIP),Beijing Municipal Science and Technology Project(Grant No.Z231100003723009)+4 种基金the Central Public-interest Scientific Institution Basal Research Fund(Grant No.Y2023PT16)TKI project(Grant No.KV1605-004)support from the China Scholarship Councilthe NPO bursary funding from Wageningen universitythe scholarship from Institute of vegetables and flowers,CAAS.
文摘Polyploidy,or whole-genome duplication,is an important evolutionary process that has shaped the genomes and traits of many plants,including numerous important crops.The Brassica genus,which includes diverse vegetables and oilseeds,is a key model system for studying how polyploidy affects plant diversification and domestication.This review summarizes the current understanding of how multiple rounds of ancient and more recent polyploidization events laid the foundation for the wide diversity seen in Brassica.We discuss the key outcomes through which polyploidy facilitates the accumulation of genetic variation,including genomic buffering that enables mutation retention.Furthermore,we explore the significant roles of interspecies and interploidy introgression in introducing external genetic novelty.We highlight homoeologous exchange(HE)as a critical mechanism unique to allopolyploids,driving substantial genomic rearrangements including presence-absence variations and gene dosage alterations that directly contribute to significant phenotypic innovation and adaptation in Brassica.Together,these polyploidy-associated processes have led to the extensive range of genomic variations that shaped great morphological diversification in the domestication of Brassica.By integrating insights from genomics,genetics,and evolutionary biology,this review shows how polyploidy has been central to Brassica's success and agricultural value.We also suggest future research areas to better understand polyploid evolution and improve crop breeding.
基金Supported by Industrial Scientific Special Project of Ministry of Agriculture(200803034)Postdoctoral Innovation Project of Shandong Province(200903017)~~
文摘[Objective] The aim was to study the feasibility of using colchicine to induce polyploidy from the buds of Shandong Salvia miltiorrhiza.[Method] The buds of Shandong Salvia miltiorrhiza were induced by colchicine.[Result] When the concentration of colchicine supplemented in culture medium was 12 mg/L,the induction rate of polyploidy was the highest（36.6%）,but its survival rate was only 16.7%.When the concentration of colchicine was 8 mg/L,the induction rate of polyploidy was 15.9%,and its survival rate was up to 40.0%.Polyploidy plant of Shandong Salvia miltiorrhiza was strong and had thick,large and dark leaves,and long stomata on the lower epidermis of leaves had small density.[Conclusion] Colchicine could induce buds effectively,and the polyploidy breeding material of Salvia miltiorrhiza was obtained,which laid a foundation for shortening breeding process.
基金supported by the Natural Science Foundation of China (31570385)
文摘Dasiphora fruticosa comprises male,female and hermaphrodite plants,which are distributed sympatrically in some populations on the Qinghai-Tibet Plateau.To explore what governs the coexistence of these three sexual phenotypes,we investigated the DNA contents,pollen and ovule production,pollen deposition,and performed hand-pollination in both hermaphroditic and dioecious individuals of D.fruticosa.Flow cytometry confirmed that the DNA content of males and females were almost twice as much as that of the hermaphrodites.Male and female flowers produced more pollen grains and ovules than hermaphroditic flowers.Hand-pollinated treatments showed that unisexual flowers were sterile in one sexual function and bisexual flowers were fertile for both functions,but no sterile seeds were produced between unisexual and bisexual flowers.Our findings imply that polyploidy is related to gender dimorphism,and both are likely to play a strong role in the coexistence of two cryptic biological species of D.fruticosa(low ploidy hermaphroditic species and high ploidy dioecious species)in the Qinghai-Tibet Plateau.
文摘Possibility of polyploidy induction by colchicine in “Balady” mandarins was investigated in vitro. Seeds were immersed in different concentrations of colchicine solutions (0.01%, 0.05%, 0.1% and 0.2%) for different durations (12, 24 and 48 hr), then cultured in vitro on MS medium at half strength. Seed survival percentage decreased by increasing colchicine concentration and duration of treatment showing the lowest percentage at 0.2% for 48 hr. The highest DNA content was recorded at 0.2% for 24 hr. Stomata No. per unit area was decreased by colchicine treatments;moreover stomata length and width were studied. The results indicated that colchicine treatment at 0.1% for 48 hr had the highest tetraploid induction efficiency percentage.
文摘Tea （Camellia sinensis L.） represents different ploidy levels. In the present paper, we reviewed the recent data on the diploid, aneuploid and polyploid formed their origin and chemically induced polyploidy and predicted it role in teagenetic improvement for better yield. Different polidy levels had different effects on tea physiology. Tetraploid and triploid had more vigour and hardness due to increased size of cells, while triploid could have even more vigours due to increased size of cells and sterilities. Chemically induced polyploidy had been found an important tool for improving plant physiology and production, therefore, induced polyploids should be produced to overcome the problem of low yield and limited rainfall in tea growing areas.
文摘An ancient genome duplication (PPP1) that predates divergence of the cereals has recently been recognized. We report here another potentially older large-scale duplication (PPP2) event that predates monocot-dicot divergence in the genome of rice (Oryza sativa L.), as inferred from the age distribution of pairs of duplicate genes based on recent genome data for rice. Our results suggest that paleopolyploidy was widespread and played an important role in the evolution of rice.
文摘This study was planned to assess genetic variation in Jatropha curcas as affected by colchichine treatments. During the experiment, seeds of Jatropha curcas were collected from mature and healthy Jatropha plants and were soaked in different concentrations of colchicines. The second treatment was that of time of soaking (seeds were soaked in different concentrations for different time durations). The results clearly revealed that by increasing the concentration of colchicine treatment, parameters like stomatal density, and size of the guard cells were reduced.
基金supported by the National Natural Science Foundation of China(Nos.32270643,91957109,and 81870427).
文摘Megakaryocytes and hepatocytes are unique cells in mammals that undergo polyploidization through endomitosis in terminal differentiation.Many polyploidization regulators and underlying mechanisms have been reported,most of which are tightly coupled with development,organogenesis,and cell differentiation.However,the nature of endomitosis,which involves successful entry into and exit from mitosis without complete cytokinesis,has not yet been fully elucidated.We highlight that endomitosis is a new cell fate in the cell cycle,and tetraploidy is a critical stage at the bifurcation of cell fate decision.This review summarizes the recent research progress in this area and provides novel insights into how cells manipulate mitosis toward endomitosis.Endomitotic cells can evade the tetraploidy restrictions and proceed to multiple rounds of the cell cycle.This knowledge not only deepens our understanding of endomitosis as a fundamental biological process but also offers new perspectives on the physiological and pathophysiological implications of polyploidization.
基金funded by Department of Science&Technology and Biotechnology,Govt.of West Bengal,India[Sanction No.565(Sanc.)/STBT-13015/15/11/2021-ST SEC]。
文摘Gerbera,a popular commercial cut flower with vibrant and striking colors has gained immense popularity in the floriculture industry.They are widely cultivated in various regions,making them available throughout the year.As a better alternative to conventional propagation methods(via seeds and rhizomes),plant tissue culture serves as way to avail large-scale,uniform,disease-free plantlets for commercial cultivation as well as to develop novel genotypes.In addition,it ensures production of healthy plantlets throughout the year in limited space.Based on the plant tissue culture techniques,the in vitro polyploidization,mutagenesis,and genetic transformation pave a path for creation of variation and eventually enhancing the ornamental traits to address the consumers’preferences and also facilitates in developing stress tolerant lines thereby minimizing the losses during cultivation,maintaining the quality of the flowers.This comprehensive review article presents an overview of the recent advancements on genetic improvement of gerbera via various cutting-edge plant tissue culture-based tools and techniques that contribute in enhancing the quality and efficiency of gerbera cultivation,meeting the demands of the floriculture industry while addressing the challenges of changing environment and resource limitations.
基金grants from the National Natural Science Foundation of China (grant numbers 31770244 and 31570224 to J.Q.)funds from the State Key Laboratory of Genetic Engineering at Fudan University.
文摘Dear Editor,Transcriptomic data have been widely employed to infer whole genome duplications (WGDs, or paleopolyploidy)(Jiao et al., 2011;Barker et al., 2016;Huang et al., 2016), through the use of large numbers of recognized paralogs with associated gene tree and Ks distributions, with the ability to date such polyploidy events.
文摘Polyploidy refers to two or more complete sets of chromosomes that are combined within a nucleus,and includes both autopolyploidy and allopolyploidy(Otto 2007).Polyploidy is a recurring theme in the evolutionary history of plants and it has been considered an important mechanism for speciation and also for adaptability to changing environments(Shimizu-Inatsugi et al.2017).In nature,about one-third of flowering plants are polyploids,and a great majority of them are allopolyploids.There are many cultivated allopolyploid species,such as cotton,Brassica and potato,the possible evolutionary history of which can be inferred with high certainty.
基金supported by the Project EVICARE (No.725229)of the European Research Council (ERC)to J.P.G.S.the ZonMw-TAS program (No.116002016)to J.P.G.S./Z.L.+2 种基金the Dutch Ministry of Economic Affairs,Agriculture and Innovation and the Netherlands CardioVascular Research Initiative (CVON)the Dutch Heart Foundation to J.P.G.S.Juntao Fang is supported by a Chinese Scholarship Council (CSC)fellowship program (No.201906210082).AV acknowledges support by the ERCAdG 787171,POLICE.
文摘Polyploid cells,which contain more than one set of chromosome pairs,are very common in nature.Polyploidy can provide cells with several potential benefits over their diploid counterparts,including an increase in cell size,contributing to organ growth and tissue homeostasis,and improving cellular robustness via increased tolerance to genomic stress and apoptotic signals.Here,we focus on why polyploidy in the cell occurs and which stress responses and molecular signals trigger cells to become polyploid.Moreover,we discuss its crucial roles in cell growth and tissue regeneration in the heart,liver,and other tissues.
基金supported by Chinese Academy of Sciences (XDB11030000)Ministry of Science and Technology of China and Ministry of Agriculture of China (2013CB127000, 2016YFA0500800, 2016ZX08009001-009, 2016ZX08005001-001)National Natural Science Foundation of China (31690092)
文摘Cottons are the most important fiber crops in the world.The cotton genus Gossypium has 52 species,including seven allotetraploid species and 45 diploids.Four species were domesticated and remain as crops under cultivation today:the New World allopolyploid species G.hirsutum and G.barbadense(2n=52),and the Old World diploid species G.arboreum and G.herbaceum(2n=26).The primary cultivated species is Upland cotton(G.hirsutum L.),which accounts for more than 90%of global cotton fiber production.
基金supported by grants from the National Natural Science Foundation of China (Grant Nos.32160051,42076117,and 41776166)Guangdong Basic and Applied Basic Research Foundation (Grant Nos.2022A1515012015,2023A1515012772)the Foreign Cultural and Educational Experts Project of the Ministry of Science and Technology (No.QNJ2021162001L)。
文摘Polyploidy is a major factor in the evolution of plants,yet we know little about the origin and evolution of polyploidy in intertidal species.This study aimed to identify the evolutionary transitions in three truemangrove species of the genus Acanthus distributed in the Indo-West Pacific region.For this purpose,we took an integrative approach that combined data on morphology,cytology,climatic niche,phylogeny,and biogeography of 493 samples from 42 geographic sites.Our results show that the Acanthus ilicifolius lineage distributed east of the Thai-Malay Peninsula possesses a tetraploid karyotype,which is morphologically distinct from that of the lineage on the west side.The haplotype networks and phylogenetic trees for the chloroplast genome and eight nuclear genes reveal that the tetraploid species has two sub-genomes,one each from A.ilicifolius and A.ebracteatus,the paternal and maternal parents,respectively.Population structure analysis also supports the hybrid speciation history of the new tetraploid species.The two sub-genomes of the tetraploid species diverged from their diploid progenitors during the Pleistocene.Environmental niche models revealed that the tetraploid species not only occupied the near-entire niche space of the diploids,but also expanded into novel environments.Our findings suggest that A.ilicifolius species distributed on the east side of the Thai-Malay Peninsula should be regarded as a new species,A.tetraploideus,which originated from hybridization between A.ilicifolius and A.ebracteatus,followed by chromosome doubling.This is the first report of a true-mangrove allopolyploid species that can reproduce sexually and clonally reproduction,which explains the long-term adaptive potential of the species.
