DAWE R K . Charting the path to fully synthetic plant chromosomes [J ] . Experimental Cell Research , 2020 , 390 ( 1 ): 111951 .

KAN M M , HUANG T B , ZHAO P P . Artificial chromosome technology and its potential application in plants [J ] . Frontiers in Plant Science , 2022 , 13 : 970943 .

BIRCHLER J A , KELLY J , SINGH J , et al . Synthetic minichromosomes in plants: past, present, and promise [J ] . The Plant Journal , 2024 , 120 ( 6 ): 2356 - 2366 .

MURRAY A W , SZOSTAK J W . Construction of artificial chromosomes in yeast [J ] . Nature , 1983 , 305 ( 5931 ): 189 - 193 .

FARR C , FANTES J , GOODFELLOW P , et al . Functional reintroduction of human telomeres into mammalian cells [J ] . Proceedings of the National Academy of Sciences of the United States of America , 1991 , 88 ( 16 ): 7006 - 7010 .

SHIZUYA H , BIRREN B , KIM U J , et al . Cloning and stable maintenance of 300-kilobase-pair fragments of human DNA in Escherichia coli using an F-factor-based vector [J ] . Proceedings of the National Academy of Sciences of the United States of America , 1992 , 89 ( 18 ): 8794 - 8797 .

HAMILTON C M , FRARY A , LEWIS C , et al . Stable transfer of intact high molecular weight DNA into plant chromosomes [J ] . Proceedings of the National Academy of Sciences of the United States of America , 1996 , 93 ( 18 ): 9975 - 9979 .

LIU Y G , SHIRANO Y , FUKAKI H , et al . Complementation of plant mutants with large genomic DNA fragments by a transformation-competent artificial chromosome vector accelerates positional cloning [J ] . Proceedings of the National Academy of Sciences of the United States of America , 1999 , 96 ( 11 ): 6535 - 6540 .

PUCHTA H , HOUBEN A . Plant chromosome engineering-past, present and future [J ] . New Phytologist , 2024 , 241 ( 2 ): 541 - 552 .

BIRCHLER J A . Engineered minichromosomes in plants [J ] . Chromosome Research , 2015 , 23 ( 1 ): 77 - 85 .

CHOO K H A . Engineering human chromosomes for gene therapy studies [J ] . Trends in Molecular Medicine , 2001 , 7 ( 6 ): 235 - 237 .

SAFFERY R , WONG L H , IRVINE D V , et al . Construction of neocentromere-based human minichromosomes by telomere-associated chromosomal truncation [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2001 , 98 ( 10 ): 5705 - 5710 .

HELLER R , BROWN K E , BURGTORF C , et al . Mini-chromosomes derived from the human Y chromosome by telomere directed chromosome breakage [J ] . Proceedings of the National Academy of Sciences of the United States of America , 1996 , 93 ( 14 ): 7125 - 7130 .

Analysis of meiotic chromosome pairing in the female mouse using a novel minichromosome [J ] . Chromosome Research , 1998 , 6 ( 4 ): 269 - 276 .

AURICHE C , DONINI P , ASCENZIONI F . Molecular and cytological analysis of a 5.5 Mb minichromosome [J ] . EMBO Reports , 2001 , 2 ( 2 ): 102 - 107 .

MORALLI D , VAGNARELLI P , BENSI M , et al . Insertion of a loxP site in a size-reduced human accessory chromosome [J ] . Cytogenetics and Cell Genetics , 2001 , 94 ( 3-4 ): 113 - 120 .

CARINE K , JACQUEMIN-SABLON A , WALTZER E , et al . Molecular characterization of human minichromosomes with centromere from chromosome 1 in human-hamster hybrid cells [J ] . Somatic Cell and Molecular Genetics , 1989 , 15 ( 5 ): 445 - 460 .

CSONKA E , CSERPÁN I , FODOR K , et al . Novel generation of human satellite DNA-based artificial chromosomes in mammalian cells [J ] . Journal of Cell Science , 2000 , 113 ( Pt 18 ): 3207 - 3216 .

LAZAR N H , CELIK S , CHEN L , et al . High-resolution genome-wide mapping of chromosome-arm-scale truncations induced by CRISPR-Cas9 editing [J ] . Nature Genetics , 2024 , 56 ( 7 ): 1482 - 1493 .

UNO N , HIRAMATSU K , UNO K , et al . CRISPR/Cas9-induced transgene insertion and telomere-associated truncation of a single human chromosome for chromosome engineering in CHO and A9 cells [J ] . Scientific Reports , 2017 , 7 : 12739 .

NAHMAD A D , REUVENI E , GOLDSCHMIDT E , et al . Frequent aneuploidy in primary human T cells after CRISPR-Cas9 cleavage [J ] . Nature Biotechnology , 2022 , 40 ( 12 ): 1807 - 1813 .

FAJKUS J , SÝKOROVÁ E , LEITCH A R . Techniques in plant telomere biology [J ] . BioTechniques , 2005 , 38 ( 2 ): 233 - 243 .

YU W C , LAMB J C , HAN F P , et al . Telomere-mediated chromosomal truncation in maize [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2006 , 103 ( 46 ): 17331 - 17336 .

YU W C , HAN F P , GAO Z , et al . Construction and behavior of engineered minichromosomes in maize [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2007 , 104 ( 21 ): 8924 - 8929 .

BIRCHLER J A , YANG H . The supernumerary B chromosome of maize: drive and genomic conflict [J ] . Open Biology , 2021 , 11 ( 11 ): 210197 .

CHEN J Y , BIRCHLER J A , HOUBEN A . The non-Mendelian behavior of plant B chromosomes [J ] . Chromosome Research , 2022 , 30 ( 2 ): 229 - 239 .

KOO D H , HAN F P , BIRCHLER J A , et al . Distinct DNA methylation patterns associated with active and inactive centromeres of the maize B chromosome [J ] . Genome Research , 2011 , 21 ( 6 ): 908 - 914 .

HUANG W , DU Y , ZHAO X , et al . B chromosome contains active genes and impacts the transcription of A chromosomes in maize ( Zea mays L.) [J ] . BMC Plant Biology , 2016 , 16 ( 1 ): 88 .

BOUDICHEVSKAIA A , FIEBIG A , KUMKE K , et al . Rye B chromosomes differently influence the expression of A chromosome-encoded genes depending on the host species [J ] . Chromosome Research , 2022 , 30 ( 4 ): 335 - 349 .

LIU X , CHEN Z Q , SHI X W . The B chromosome: an optimum platform for maize minichromosome engineering [J ] . Critical Reviews in Plant Sciences , 2025 , 44 ( 1 ): 30 - 45 .

NELSON A D , LAMB J C , KOBROSSLY P S , et al . Parameters affecting telomere-mediated chromosomal truncation in Arabidopsis [J ] . The Plant Cell , 2011 , 23 ( 6 ): 2263 - 2272 .

TEO C H , MA L , KAPUSI E , et al . Induction of telomere-mediated chromosomal truncation and stability of truncated chromosomes in Arabidopsis thaliana [J ] . The Plant Journal , 2011 , 68 ( 1 ): 28 - 39 .

MURATA M . Artificial chromosome preparation in Arabidopsis [J ] . Current Protocols in Plant Biology , 2016 , 1 ( 1 ): 53 - 66 .

XU C H , CHENG Z K , YU W C . Construction of rice mini-chromosomes by telomere-mediated chromosomal truncation [J ] . The Plant Journal , 2012 , 70 ( 6 ): 1070 - 1079 .

YANG X Y , LI J H , CHEN L , et al . Stable mitotic inheritance of rice minichromosomes in cell suspension cultures [J ] . Plant Cell Reports , 2015 , 34 ( 6 ): 929 - 941 .

KAPUSI E , MA L , TEO C H , et al . Telomere-mediated truncation of barley chromosomes [J ] . Chromosoma , 2012 , 121 ( 2 ): 181 - 190 .

YUAN J , SHI Q H , GUO X , et al . Site-specific transfer of chromosomal segments and genes in wheat engineered chromosomes [J ] . Journal of Genetics and Genomics , 2017 , 44 ( 11 ): 531 - 539 .

YAN X H , LI C , YANG J , et al . Induction of telomere-mediated chromosomal truncation and behavior of truncated chromosomes in Brassica napus [J ] . The Plant Journal , 2017 , 91 ( 4 ): 700 - 713 .

YIN X Z , ZHANG Y X , CHEN Y H , et al . Precise characterization and tracking of stably inherited artificial minichromosomes made by telomere-mediated chromosome truncation in Brassica napus [J ] . Frontiers in Plant Science , 2021 , 12 : 743792 .

LIU Y , LIU Q , YI C Y , et al . Past innovations and future possibilities in plant chromosome engineering [J ] . Plant Biotechnology Journal , 2025 , 23 ( 3 ): 695 - 708 .

SEARS E R . The transfer of leaf-rust resistance from Aegilops umbellulata to wheat [J ] . Genetics in plant breeding. Brook-haven Symposia in Biology , 1956 , 9 : 1 - 22 .

FU S L , LV Z L , QI B , et al . Molecular cytogenetic characterization of wheat: Thinopyrum elongatum addition, substitution and translocation lines with a novel source of resistance to wheat Fusarium head blight [J ] . Journal of Genetics and Genomics , 2012 , 39 ( 2 ): 103 - 110 .

GUO X R , SHI Q H , LIU Y , et al . Systemic development of wheat- Thinopyrum elongatum translocation lines and their deployment in wheat breeding for Fusarium head blight resistance [J ] . The Plant Journal , 2023 , 114 ( 6 ): 1475 - 1489 .

KASZÁS E , BIRCHLER J A . Meiotic transmission rates correlate with physical features of rearranged centromeres in maize [J ] . Genetics , 1998 , 150 ( 4 ): 1683 - 1692 .

YAMADA T , FUJIMOTO Y , YAMAMOTO Y , et al . Minichromosome formation in Chlorella cells irradiated with electron beams [J ] . Journal of Bioscience and Bioengineering , 2003 , 95 ( 6 ): 601 - 607 .

LYZNIK L A , GORDON-KAMM W J , TAO Y . Site-specific recombination for genetic engineering in plants [J ] . Plant Cell Reports , 2003 , 21 ( 10 ): 925 - 932 .

SUN C , LEI Y , LI B S , et al . Precise integration of large DNA sequences in plant genomes using PrimeRoot editors [J ] . Nature Biotechnology , 2024 , 42 ( 2 ): 316 - 327 .

MURATA M , SHIBATA F , HIRONAKA A , et al . Generation of an artificial ring chromosome in Arabidopsis by Cre/LoxP-mediated recombination [J ] . The Plant Journal , 2013 , 74 ( 3 ): 363 - 371 .

MURATA M . Minichromosomes and artificial chromosomes in Arabidopsis [J ] . Chromosome Research , 2014 , 22 ( 2 ): 167 - 178 .

GAETA R T , MASONBRINK R E , ZHAO C Z , et al . In vivo modification of a maize engineered minichromosome [J ] . Chromosoma , 2013 , 122 ( 3 ): 221 - 232 .

BARRANGOU R , FREMAUX C , DEVEAU H , et al . CRISPR provides acquired resistance against viruses in prokaryotes [J ] . Science , 2007 , 315 ( 5819 ): 1709 - 1712 .

JINEK M , CHYLINSKI K , FONFARA I , et al . A programmable dual-RNA-guided DNA endonuclease in adaptive bacterial immunity [J ] . Science , 2012 , 337 ( 6096 ): 816 - 821 .

ZHOU H B , LIU B , WEEKS D P , et al . Large chromosomal deletions and heritable small genetic changes induced by CRISPR/Cas9 in rice [J ] . Nucleic Acids Research , 2014 , 42 ( 17 ): 10903 - 10914 .

DURR J , PAPAREDDY R , NAKAJIMA K , et al . Highly efficient heritable targeted deletions of gene clusters and non-coding regulatory regions in Arabidopsis using CRISPR/Cas9 [J ] . Scientific Reports , 2018 , 8 : 4443 .

LI R Q , CHAR S N , YANG B . Creating large chromosomal deletions in rice using CRISPR/Cas9 [M/OL ] //QI Y. Plant genome editing with CRISPR systems . New York: Springer New York, 2019 : 47 - 61 . ( 2019-01-05)[2025-09-01] . https://doi.org/10.1007/978-1-4939-8991-1_4 https://doi.org/10.1007/978-1-4939-8991-1_4 .

LI Y N , HUANG B Y , CHEN J , et al . Targeted large fragment deletion in plants using paired crRNAs with type Ⅰ CRISPR system [J ] . Plant Biotechnology Journal , 2023 , 21 ( 11 ): 2196 - 2208 .

RÖNSPIES M , SCHMIDT C , SCHINDELE P , et al . Massive crossover suppression by CRISPR-Cas-mediated plant chromosome engineering [J ] . Nature Plants , 2022 , 8 ( 10 ): 1153 - 1159 .

SCHMIDT C , PACHER M , PUCHTA H . Efficient induction of heritable inversions in plant genomes using the CRISPR/Cas system [J ] . The Plant Journal , 2019 , 98 ( 4 ): 577 - 589 .

SCHWARTZ C , LENDERTS B , FEIGENBUTZ L , et al . CRISPR-Cas9-mediated 75.5-Mb inversion in maize [J ] . Nature Plants , 2020 , 6 ( 12 ): 1427 - 1431 .

MAO Y H , ZHAO Y L , ZHOU Q , et al . Chromosome engineering: technologies, applications, and challenges [J ] . Annual Review of Animal Biosciences , 2025 , 13 ( 1 ): 25 - 47 .

WANG M L , LIN X J , MO B X , et al . Plant artificial chromosomes: construction and transformation [J ] . ACS Synthetic Biology , 2024 , 13 ( 1 ): 15 - 24 .

SHIZUYA H , KOUROS-MEHR H . The development and applications of the bacterial artificial chromosome cloning system [J ] . The Keio Journal of Medicine , 2001 , 50 ( 1 ): 26 - 30 .

HAO M L , TANG J B , GE S X , et al . Bacterial-artificial-chromosome-based genome editing methods and the applications in herpesvirus research [J ] . Microorganisms , 2023 , 11 ( 3 ): 589 .

KUDO K , NISHIMURA T , IZUMIKAWA M , et al . Capability of a large bacterial artificial chromosome clone harboring multiple biosynthetic gene clusters for the production of diverse compounds [J ] . The Journal of Antibiotics , 2024 , 77 ( 5 ): 288 - 298 .

MOGI K , TOMITA H , YOSHIHARA M , et al . Advances in bacterial artificial chromosome (BAC) transgenic mice for gene analysis and disease research [J ] . Gene , 2025 , 934 : 149014 .

CARLSON S R , RUDGERS G W , ZIELER H , et al . Meiotic transmission of an in vitro -assembled autonomous maize minichromosome [J ] . PLoS Genetics , 2007 , 3 ( 10 ): e179 .

ANANIEV E V , WU C C , CHAMBERLIN M A , et al . Artificial chromosome formation in maize ( Zea mays L.) [J ] . Chromosoma , 2009 , 118 ( 2 ): 157 - 177 .

HAMILTON C M . A binary-BAC system for plant transformation with high-molecular-weight DNA [J ] . Gene , 1997 , 200 ( 1-2 ): 107 - 116 .

SHIBATA D , SEKI M , MITSUKAWA N , et al . Establishment of framework P1 clones for map-based cloning and genome sequencing: direct RFLP mapping of large clones [J ] . Gene , 1998 , 225 ( 1-2 ): 31 - 38 .

MOZO T , DEWAR K , DUNN P , et al . A complete BAC-based physical map of the Arabidopsis thaliana genome [J ] . Nature Genetics , 1999 , 22 ( 3 ): 271 - 275 .

LIU Y G , NAGAKI K , FUJITA M , et al . Development of an efficient maintenance and screening system for large-insert genomic DNA libraries of hexaploid wheat in a transformation-competent artificial chromosome (TAC) vector [J ] . The Plant Journal , 2000 , 23 ( 5 ): 687 - 695 .

LIU Y G , LIU H M , CHEN L T , et al . Development of new transformation-competent artificial chromosome vectors and rice genomic libraries for efficient gene cloning [J ] . Gene , 2002 , 282 ( 1-2 ): 247 - 255 .

HIROSE Y , SUDA K , LIU Y G , et al . The Arabidopsis TAC Position Viewer: a high-resolution map of transformation-competent artificial chromosome (TAC) clones aligned with the Arabidopsis thaliana Columbia-0 genome [J ] . The Plant Journal , 2015 , 83 ( 6 ): 1114 - 1122 .

QU S , COAKER G , FRANCIS D , et al . Development of a new transformation-competent artificial chromosome (TAC) vector and construction of tomato and rice TAC libraries [J ] . Molecular Breeding , 2003 , 12 ( 4 ): 297 - 308 .

LIANG F S , ZHANG K C , YU Z W , et al . Construction, characterization, and screening of a transformation-competent artificial chromosome library of peach [J ] . Plant Molecular Biology Reporter , 2004 , 22 ( 1 ): 37 - 48 .

KONG F N , JIANG S M , SHI L X , et al . Construction and characterization of a transformation-competent artificial chromosome (TAC) library of Zizania latifolia (Griseb.) [J ] . Plant Molecular Biology Reporter , 2006 , 24 ( 2 ): 219 - 227 .

WANG J Y , XIE Z X , CUI Y Z , et al . Artificial design of the genome: from sequences to the 3D structure of chromosomes [J ] . Trends in Biotechnology , 2025 , 43 ( 2 ): 304 - 317 .

CELLO J , PAUL A V , WIMMER E . Chemical synthesis of poliovirus cDNA: generation of infectious virus in the absence of natural template [J ] . Science , 2002 , 297 ( 5583 ): 1016 - 1018 .

SMITH H O , HUTCHISON C A , PFANNKOCH C , et al . Generating a synthetic genome by whole genome assembly: ϕX174 bacteriophage from synthetic oligonucleotides [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2003 , 100 ( 26 ): 15440 - 15445 .

RICHARDSON S M , MITCHELL L A , STRACQUADANIO G , et al . Design of a synthetic yeast genome [J ] . Science , 2017 , 355 ( 6329 ): 1040 - 1044 .

THI NHU THAO T , LABROUSSAA F , EBERT N , et al . Rapid reconstruction of SARS-CoV-2 using a synthetic genomics platform [J ] . Nature , 2020 , 582 ( 7813 ): 561 - 565 .

CHEN L G , LAN T L , ZHANG S , et al . A designer synthetic chromosome fragment functions in moss [J ] . Nature Plants , 2024 , 10 ( 2 ): 228 - 239 .

朱骊宇 , 赵玉龙 , 李伟 , 等 . 哺乳动物染色体工程研究进展 [J ] . 合成生物学 , 2023 , 4 ( 2 ): 394 - 406 .

ZHU L Y , ZHAO Y L , LI W , et al . Progress in mammalian chromosome engineering [J ] . Synthetic Biology Journal , 2023 , 4 ( 2 ): 394 - 406 .

ELENA C , RAVASI P , CASTELLI M E , et al . Expression of codon optimized genes in microbial systems: current industrial applications and perspectives [J ] . Frontiers in Microbiology , 2014 , 5 : 21 .

OSTROV N , LANDON M , GUELL M , et al . Design, synthesis, and testing toward a 57-codon genome [J ] . Science , 2016 , 353 ( 6301 ): 819 - 822 .

HOCHREIN L , MITCHELL L A , SCHULZ K , et al . L-SCRaMbLE as a tool for light-controlled Cre-mediated recombination in yeast [J ] . Nature Communications , 2018 , 9 : 1931 .

DYMOND J S , RICHARDSON S M , COOMBES C E , et al . Synthetic chromosome arms function in yeast and generate phenotypic diversity by design [J ] . Nature , 2011 , 477 ( 7365 ): 471 - 476 .

ANNALURU N , MULLER H , MITCHELL L A , et al . Total synthesis of a functional designer eukaryotic chromosome [J ] . Science , 2014 , 344 ( 6179 ): 55 - 58 .

SHEN Y , WANG Y , CHEN T , et al . Deep functional analysis of synⅡ, a 770-kilobase synthetic yeast chromosome [J ] . Science , 2017 , 355 ( 6329 ): eaaf4791 .

XIE Z X , LI B Z , MITCHELL L A , et al . “Perfect” designer chromosome Ⅴ and behavior of a ring derivative [J ] . Science , 2017 , 355 ( 6329 ): eaaf4704 .

MITCHELL L A , WANG A , STRACQUADANIO G , et al . Synthesis, debugging, and effects of synthetic chromosome consolidation: synⅥ and beyond [J ] . Science , 2017 , 355 ( 6329 ): eaaf4831 .

WU Y , LI B Z , ZHAO M , et al . Bug mapping and fitness testing of chemically synthesized chromosome X [J ] . Science , 2017 , 355 ( 6329 ): eaaf4706 .

SCHINDLER D , WALKER R S K , JIANG S Y , et al . Design, construction, and functional characterization of a tRNA neochromosome in yeast [J ] . Cell , 2023 , 186 ( 24 ): 5237 - 5253.e22 .

DU F , DAI J B , JIAO Y L . Insights into a functional synthetic plant genome [J ] . New Phytologist , 2024 , 244 ( 1 ): 46 - 50 .

YE H , LUO G Y , ZHENG Z W , et al . Plant synthetic genomics: big lessons from the little yeast [J ] . Cell Chemical Biology , 2024 , 31 ( 10 ): 1745 - 1754 .

YU W F , ZHANG S , ZHAO S J , et al . Designing a synthetic moss genome using GenoDesigner [J ] . Nature Plants , 2024 , 10 ( 6 ): 848 - 856 .

LANG D , ULLRICH K K , MURAT F , et al . The Physcomitrella patens chromosome-scale assembly reveals moss genome structure and evolution [J ] . The Plant Journal , 2018 , 93 ( 3 ): 515 - 533 .

BI G Q , ZHAO S J , YAO J W , et al . Near telomere-to-telomere genome of the model plant Physcomitrium patens [J ] . Nature Plants , 2024 , 10 ( 2 ): 327 - 343 .

ISAACS F J , CARR P A , WANG H H , et al . Precise manipulation of chromosomes in vivo enables genome-wide codon replacement [J ] . Science , 2011 , 333 ( 6040 ): 348 - 353 .

FREDENS J , WANG K H , DE LA TORRE D , et al . Total synthesis of Escherichia coli with a recoded genome [J ] . Nature , 2019 , 569 ( 7757 ): 514 - 518 .

ROVNER A J , HAIMOVICH A D , KATZ S R , et al . Recoded organisms engineered to depend on synthetic amino acids [J ] . Nature , 2015 , 518 ( 7537 ): 89 - 93 .

CHENG L , ZHAO S J , LI T Y , et al . Large-scale genomic rearrangements boost SCRaMbLE in Saccharomyces cerevisiae [J ] . Nature Communications , 2024 , 15 : 770 .

BLOUNT B A , GOWERS G F , HO J C H , et al . Rapid host strain improvement by in vivo rearrangement of a synthetic yeast chromosome [J ] . Nature Communications , 2018 , 9 : 1932 .

LUO Z Q , WANG L H , WANG Y , et al . Identifying and characterizing SCRaMbLEd synthetic yeast using ReSCuES [J ] . Nature Communications , 2018 , 9 : 1930 .

JIA B , WU Y , LI B Z , et al . Precise control of SCRaMbLE in synthetic haploid and diploid yeast [J ] . Nature Communications , 2018 , 9 : 1933 .

SCHINDLER D , DAI J B , CAI Y Z . Synthetic genomics: a new venture to dissect genome fundamentals and engineer new functions [J ] . Current Opinion in Chemical Biology , 2018 , 46 : 56 - 62 .

JIANG S Y , TANG Y W , XIANG L , et al . Efficient de novo assembly and modification of large DNA fragments [J ] . Science China Life Sciences , 2022 , 65 ( 7 ): 1445 - 1455 .

SHAO Y Y , LU N , WU Z F , et al . Creating a functional single-chromosome yeast [J ] . Nature , 2018 , 560 ( 7718 ): 331 - 335 .

GU X , YE T T , ZHANG X R , et al . Single-chromosome fission yeast models reveal the configuration robustness of a functional genome [J ] . Cell Reports , 2022 , 40 ( 8 ): 111237 .

WANG L B , LI Z K , WANG L Y , et al . A sustainable mouse karyotype created by programmed chromosome fusion [J ] . Science , 2022 , 377 ( 6609 ): 967 - 975 .

MERCY G , MOZZICONACCI J , SCOLARI V F , et al . 3D organization of synthetic and scrambled chromosomes [J ] . Science , 2017 , 355 ( 6329 ): eaaf4597 .

ZHANG W M , LAZAR-STEFANITA L , YAMASHITA H , et al . Manipulating the 3D organization of the largest synthetic yeast chromosome [J ] . Molecular Cell , 2023 , 83 ( 23 ): 4424 - 4437.e5 .

WESTHORPE F G , STRAIGHT A F . Chromosome segregation: reconstituting the kinetochore [J ] . Current Biology , 2016 , 26 ( 23 ): R1242 - R1245 .

EVATT J M , SADLI A D , RAPACZ B K , et al . Centromere pairing enables correct segregation of meiotic chromosomes [J ] . Current Biology , 2024 , 34 ( 10 ): 2085 - 2093.e6 .

CLEVELAND D W , MAO Y H , SULLIVAN K F . Centromeres and kinetochores: from epigenetics to mitotic checkpoint signaling [J ] . Cell , 2003 , 112 ( 4 ): 407 - 421 .

PROSÉE R F , WENDA J M , STEINER F A . Adaptations for centromere function in meiosis [J ] . Essays in Biochemistry , 2020 , 64 ( 2 ): 193 - 203 .

NAISH M , HENDERSON I R . The structure, function, and evolution of plant centromeres [J ] . Genome Research , 2024 , 34 ( 2 ): 161 - 178 .

KOCH L B , MARSTON A L . The functional organisation of the centromere and kinetochore during meiosis [J ] . Current Opinion in Cell Biology , 2025 , 94 : 102486 .

MELTERS D P , BRADNAM K R , YOUNG H A , et al . Comparative analysis of tandem repeats from hundreds of species reveals unique insights into centromere evolution [J ] . Genome Biology , 2013 , 14 ( 1 ): R10 .

WLODZIMIERZ P , RABANAL F A , BURNS R , et al . Cycles of satellite and transposon evolution in Arabidopsis centromeres [J ] . Nature , 2023 , 618 ( 7965 ): 557 - 565 .

LOGSDON G A , ROZANSKI A N , RYABOV F , et al . The variation and evolution of complete human centromeres [J ] . Nature , 2024 , 629 ( 8010 ): 136 - 145 .

COPENHAVER G P , NICKEL K , KUROMORI T , et al . Genetic definition and sequence analysis of Arabidopsis centromeres [J ] . Science , 1999 , 286 ( 5449 ): 2468 - 2474 .

NAGAKI K , CHENG Z K , OUYANG S , et al . Sequencing of a rice centromere uncovers active genes [J ] . Nature Genetics , 2004 , 36 ( 2 ): 138 - 145 .

NAISH M , ALONGE M , WLODZIMIERZ P , et al . The genetic and epigenetic landscape of the Arabidopsis centromeres [J ] . Science , 2021 , 374 ( 6569 ): eabi7489 .

CHENG Z K , DONG F G , LANGDON T , et al . Functional rice centromeres are marked by a satellite repeat and a centromere-specific retrotransposon [J ] . The Plant Cell , 2002 , 14 ( 8 ): 1691 - 1704 .

ZHONG C X , MARSHALL J B , TOPP C , et al . Centromeric retroelements and satellites interact with maize kinetochore protein CENH3 [J ] . The Plant Cell , 2002 , 14 ( 11 ): 2825 - 2836 .

NAGAKI K , NEUMANN P , ZHANG D F , et al . Structure, divergence, and distribution of the CRR centromeric retrotransposon family in rice [J ] . Molecular Biology and Evolution , 2005 , 22 ( 4 ): 845 - 855 .

SONG J M , XIE W Z , WANG S , et al . Two gap-free reference genomes and a global view of the centromere architecture in rice [J ] . Molecular Plant , 2021 , 14 ( 10 ): 1757 - 1767 .

CHEN J , WANG Z J , TAN K W , et al . A complete telomere-to-telomere assembly of the maize genome [J ] . Nature Genetics , 2023 , 55 ( 7 ): 1221 - 1231 .

CLARKE L , CARBON J . Isolation of a yeast centromere and construction of functional small circular chromosomes [J ] . Nature , 1980 , 287 ( 5782 ): 504 - 509 .

KOUPRINA N , EBERSOLE T , KORIABINE M , et al . Cloning of human centromeres by transformation-associated recombination in yeast and generation of functional human artificial chromosomes [J ] . Nucleic Acids Research , 2003 , 31 ( 3 ): 922 - 934 .

TALBERT P B , MASUELLI R , TYAGI A P , et al . Centromeric localization and adaptive evolution o f an Arabidopsis histone H3 variant [J ] . The Plant Cell , 2002 , 14 ( 5 ): 1053 - 1066 .

HAN F P , LAMB J C , BIRCHLER J A . High frequency of centromere inactivation resulting in stable dicentric chromosomes of maize [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2006 , 103 ( 9 ): 3238 - 3243 .

PHAN B H , JIN W W , TOPP C N , et al . Transformation of rice with long DNA-segments consisting of random genomic DNA or centromere-specific DNA [J ] . Transgenic Research , 2007 , 16 ( 3 ): 341 - 351 .

HARA M , FUKAGAWA T . Where is the right path heading from the centromere to spindle microtubules? [J ] . Cell Cycle , 2019 , 18 ( 11 ): 1199 - 1211 .

RAIPURIA R K , WATTS A , SHARMA B B , et al . Decoding allelic diversity, transcript variants and transcriptional complexity of CENH3 gene in Brassica oleracea var. botrytis [J ] . Protoplasma , 2023 , 260 ( 4 ): 1149 - 1162 .

LERMONTOVA I , SCHUBERT V , FUCHS J , et al . Loading of Arabidopsis centromeric histone CENH3 occurs mainly during G2 and requires the presence of the histone fold domain [J ] . The Plant Cell , 2006 , 18 ( 10 ): 2443 - 2451 .

LERMONTOVA I , KOROLEVA O , RUTTEN T , et al . Knockdown of CENH3 in Arabidopsis reduces mitotic divisions and causes sterility by disturbed meiotic chromosome segregation [J ] . The Plant Journal , 2011 , 68 ( 1 ): 40 - 50 .

KARIMI-ASHTIYANI R , ISHII T , NIESSEN M , et al . Point mutation impairs centromeric CENH3 loading and induces haploid plants [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2015 , 112 ( 36 ): 11211 - 11216 .

RAVI M , CHAN S W L . Haploid plants produced by centromere-mediated genome elimination [J ] . Nature , 2010 , 464 ( 7288 ): 615 - 618 .

KELLIHER T , STARR D , WANG W L , et al . Maternal haploids are preferentially induced by CENH3-tailswap transgenic complementation in maize [J ] . Frontiers in Plant Science , 2016 , 7 : 414 .

RAVI M , SHIBATA F , RAMAHI J S , et al . Meiosis-specific loading of the centromere-specific histone CENH3 in Arabidopsis thaliana [J ] . PLoS Genetics , 2011 , 7 ( 6 ): e1002121 .

WANG N , DAWE R K . Centromere size and its relationship to haploid formation in plants [J ] . Molecular Plant , 2018 , 11 ( 3 ): 398 - 406 .

DUNLEAVY E M , ROCHE D , TAGAMI H , et al . HJURP is a cell-cycle-dependent maintenance and deposition factor of CENP-A at centromeres [J ] . Cell , 2009 , 137 ( 3 ): 485 - 497 .

FOLTZ D R , JANSEN L E T , BAILEY A O , et al . Centromere-specific assembly of CENP-A nucleosomes is mediated by HJURP [J ] . Cell , 2009 , 137 ( 3 ): 472 - 484 .

MENDIBURO M J , PADEKEN J , FÜLÖP S , et al . Drosophila CENH3 is sufficient for centromere formation [J ] . Science , 2011 , 334 ( 6056 ): 686 - 690 .

LOGSDON G A , GAMBOGI C W , LISKOVYKH M A , et al . Human artificial chromosomes that bypass centromeric DNA [J ] . Cell , 2019 , 178 ( 3 ): 624 - 639.e19 .

GAMBOGI C W , BIRCHAK G J , MER E , et al . Efficient formation of single-copy human artificial chromosomes [J ] . Science , 2024 , 383 ( 6689 ): 1344 - 1349 .

TEO C H , LERMONTOVA I , HOUBEN A , et al . De novo generation of plant centromeres at tandem repeats [J ] . Chromosoma , 2013 , 122 ( 3 ): 233 - 241 .

DAWE R K , GENT J I , ZENG Y B , et al . Synthetic maize centromeres transmit chromosomes across generations [J ] . Nature Plants , 2023 , 9 ( 3 ): 433 - 441 .

ZENG Y B , WANG M Y , GENT J I , et al . Increased maize chromosome number by engineered chromosome fission [J ] . Science Advances , 2025 , 11 ( 21 ): eadw3433 .

BAI S , LUO H , TONG H Z , et al . Advances on transfer and maintenance of large DNA in bacteria, fungi, and mammalian cells [J ] . Biotechnology Advances , 2024 , 76 : 108421 .

CAMPELO S N , HUANG P H , BUIE C R , et al . Recent advancements in electroporation technologies: from bench to clinic [J ] . Annual Review of Biomedical Engineering , 2023 , 25 : 77 - 100 .

LEE E C , LIANG Q , ALI H , et al . Complete humanization of the mouse immunoglobulin loci enables efficient therapeutic antibody discovery [J ] . Nature Biotechnology , 2014 , 32 ( 4 ): 356 - 363 .

MITCHELL L A , MCCULLOCH L H , PINGLAY S , et al . De novo assembly and delivery to mouse cells of a 101 kb functional human gene [J ] . Genetics , 2021 , 218 ( 1 ): iyab038 .

MARSCHALL P , MALIK N , LARIN Z . Transfer of YACs up to 2.3 Mb intact into human cells with polyethylenimine [J ] . Gene Therapy , 1999 , 6 ( 9 ): 1634 - 1637 .

MEJÍA J E , WILLMOTT A , LEVY E , et al . Functional complementation of a genetic deficiency with human artificial chromosomes [J ] . The American Journal of Human Genetics , 2001 , 69 ( 2 ): 315 - 326 .

MARTINEZ A , KOLVEK S J , YIP C L T , et al . Genetically modified bacterial strains and novel bacterial artificial chromosome shuttle vectors for constructing environmental libraries and detecting heterologous natural products in multiple expression hosts [J ] . Applied and Environmental Microbiology , 2004 , 70 ( 4 ): 2452 - 2463 .

BAÑUELOS-VAZQUEZ L A , TORRES TEJERIZO G , BROM S . Regulation of conjugative transfer of plasmids and integrative conjugative elements [J ] . Plasmid , 2017 , 91 : 82 - 89 .

BROPHY J A N , TRIASSI A J , ADAMS B L , et al . Engineered integrative and conjugative elements for efficient and inducible DNA transfer to undomesticated bacteria [J ] . Nature Microbiology , 2018 , 3 ( 9 ): 1043 - 1053 .

LI L P , BLANKENSTEIN T . Generation of transgenic mice with megabase-sized human yeast artificial chromosomes by yeast spheroplast-embryonic stem cell fusion [J ] . Nature Protocols , 2013 , 8 ( 8 ): 1567 - 1582 .

BROWN D M , CHAN Y A , DESAI P J , et al . Efficient size-independent chromosome delivery from yeast to cultured cell lines [J ] . Nucleic Acids Research , 2017 , 45 ( 7 ): e50 .

EGE T , RINGERTZ N R . Preparation of microcells by enucleation of micronucleate cells [J ] . Experimental Cell Research , 1974 , 87 ( 2 ): 378 - 382 .

O’DOHERTY A , RUF S , MULLIGAN C , et al . An aneuploid mouse strain carrying human chromosome 21 with down syndrome phenotypes [J ] . Science , 2005 , 309 ( 5743 ): 2033 - 2037 .

KATOH M , KAZUKI Y , KAZUKI K , et al . Exploitation of the interaction of measles virus fusogenic envelope proteins with the surface receptor CD46 on human cells for microcell-mediated chromosome transfer [J ] . BMC Biotechnology , 2010 , 10 : 37 .

LISKOVYKH M , LEE N C O , LARIONOV V , et al . Moving toward a higher efficiency of microcell-mediated chromosome transfer [J ] . Molecular Therapy - Methods & Clinical Development , 2016 , 3 : 16043 .

MIYAMOTO H , KOBAYASHI H , KISHIMA N , et al . Rapid human genomic DNA cloning into mouse artificial chromosome via direct chromosome transfer from human iPSC and CRISPR/Cas9-mediated translocation [J ] . Nucleic Acids Research , 2024 , 52 ( 3 ): 1498 - 1511 .

WANG Y F , ZENG H Y , ZHOU X , et al . Transformation of rice with large maize genomic DNA fragments containing high content repetitive sequences [J ] . Plant Cell Reports , 2015 , 34 ( 6 ): 1049 - 1061 .

MULLEN J , ADAM G , BLOWERS A , et al . Biolistic transfer of large DNA fragments to tobacco cells using YACs retrofitted for plant transformation [J ] . Molecular Breeding , 1998 , 4 ( 5 ): 449 - 457 .

ZHAO X , MENG Z G , WANG Y , et al . Pollen magnetofection for genetic modification with magnetic nanoparticles as gene carriers [J ] . Nature Plants , 2017 , 3 ( 12 ): 956 - 964 .

VAN WORDRAGEN M , SHAKYA R , VERKERK R , et al . Liposome-mediated transfer of YAC-DNA to tobacco cells [J ] . Plant Molecular Biology Reporter , 1997 , 15 ( 2 ): 170 - 178 .

逄春梅 , 张亮 , 孙春玉 , 等 . 人参大片段DNA(100kb)转化灵芝的研究 [J ] . 菌物学报 , 2013 , 32 ( 1 ): 96 - 102 .

PANG C M , ZHANG L , SUN C Y , et al . Transformation of large DNA fragment of Panax ginseng into Ganoderma lucidum [J ] . Mycosystema , 2013 , 32 ( 1 ): 96 - 102 .

HAMILTON C M , FRARY A , XU Y M , et al . Construction of tomato genomic DNA libraries in a binary-BAC (BIBAC) vector [J ] . The Plant Journal , 1999 , 18 ( 2 ): 223 - 229 .

SHIBATA D , LIU Y G . Technical focus- Agrobacterium -mediated plant transformation with large DNA fragments [J ] . Trends in Plant Science , 2000 , 5 ( 8 ): 354 - 357 .

VEGA J M , YU W C , HAN F P , et al . Agrobacterium -mediated transformation of maize ( Zea mays ) with Cre-lox site specific recombination cassettes in BIBAC vectors [J ] . Plant Molecular Biology , 2008 , 66 ( 6 ): 587 - 598 .

HE R F , WANG Y Y , SHI Z Y , et al . Construction of a genomic library of wild rice and Agrobacterium -mediated transformation of large insert DNA linked to BPH resistance locus [J ] . Gene , 2003 , 321 : 113 - 121 .

HE R F , PAN J , ZHU L L , et al . Agrobacterium -mediated transformation of large DNA fragments using a BIBAC vector system in rice [J ] . Plant Molecular Biology Reporter , 2010 , 28 ( 4 ): 613 - 619 .

KLEIN T M , WOLF E D , WU R , et al . High-velocity microprojectiles for delivering nucleic acids into living cells [J ] . Nature , 1987 , 327 ( 6117 ): 70 - 73 .

ALTPETER F , BAISAKH N , BEACHY R , et al . Particle bombardment and the genetic enhancement of crops: myths and realities [J ] . Molecular Breeding , 2005 , 15 ( 3 ): 305 - 327 .

LACROIX B , CITOVSKY V . Biolistic approach for transient gene expression studies in plants [M/OL ] //RUSTGI S, LUO H. Methods in molecular biology: biolistic DNA delivery in plants . New York: Springer US, 2020 : 125 - 139 . ( 2020-04-11)[2025-09-01] . https://doi.org/10.1007/978-1-0716-0356-7_6 https://doi.org/10.1007/978-1-0716-0356-7_6 .

ERCOLANO M R , BALLVORA A , PAAL J , et al . Functional complementation analysis in potato via biolistic transformation with BAC large DNA fragments [J ] . Molecular Breeding , 2004 , 13 ( 1 ): 15 - 22 .

CHANG Y L , CHUANG H W , MEKSEM K , et al . Characterization of a plant-transformation-ready large-insert BIBAC library of Arabidopsis and bombardment transformation of a large-insert BIBAC of the library into tobacco [J ] . Genome , 2011 , 54 ( 6 ): 437 - 447 .

CUNNINGHAM F J , DEMIRER G S , GOH N S , et al . Nanobiolistics: an emerging genetic transformation approach [M/OL ] //RUSTGI S, LUO H. Methods in molecular biology: biolistic DNA delivery in plants . New York: Springer US, 2020 : 141 - 159 . ( 2020-04-11)[2025-09-01] . https://doi.org/10.1007/978-1-0716-0356-7_7 https://doi.org/10.1007/978-1-0716-0356-7_7 .

LV Z Y , JIANG R , CHEN J F , et al . Nanoparticle-mediated gene transformation strategies for plant genetic engineering [J ] . The Plant Journal , 2020 , 104 ( 4 ): 880 - 891 .

DEMIRER G S , ZHANG H , GOH N S , et al . Carbon nanotube-mediated DNA delivery without transgene integration in intact plants [J ] . Nature Protocols , 2019 , 14 ( 10 ): 2954 - 2971 .

MORGAN J M , JELENSKA J , HENSLEY D , et al . An efficient and broadly applicable method for transient transformation of plants using vertically aligned carbon nanofiber arrays [J ] . Frontiers in Plant Science , 2022 , 13 : 1051340 .

TORNEY F , TREWYN B G , LIN V S Y , et al . Mesoporous silica nanoparticles deliver DNA and chemicals into plants [J ] . Nature Nanotechnology , 2007 , 2 ( 5 ): 295 - 300 .

CUNNINGHAM F J , GOH N S , DEMIRER G S , et al . Nanoparticle-mediated delivery towards advancing plant genetic engineering [J ] . Trends in Biotechnology , 2018 , 36 ( 9 ): 882 - 897 .

TERAMURA Y , KANEDA Y , TOTANI T , et al . Behavior of synthetic polymers immobilized on a cell membrane [J ] . Biomaterials , 2008 , 29 ( 10 ): 1345 - 1355 .

ZHENG H Z , LIU H H , CHEN S X , et al . Yeast transformation process studied by fluorescence labeling technique [J ] . Bioconjugate Chemistry , 2005 , 16 ( 2 ): 250 - 254 .

HASEZAWA S , MATSUI C , NAGATA T , et al . Cytological study of the introduction of Agrobacterium tumefaciens spheroplasts into Vinca rosea protoplasts [J ] . Canadian Journal of Botany , 1983 , 61 ( 4 ): 1052 - 1057 .

HATSUYAMA Y , SUNAGA N , HABU Y , et al . Direct transfer of plasmid DNA from intact yeast spheroplasts into plant protoplasts [J ] . Plant & Cell Physiology , 1994 , 35 ( 1 ): 93 - 98 .

YADAV T , QUIVY J P , ALMOUZNI G . Chromatin plasticity: a versatile landscape that underlies cell fate and identity [J ] . Science , 2018 , 361 ( 6409 ): 1332 - 1336 .

BARNHART M C , KUICH P H J L , STELLFOX M E , et al . HJURP is a CENP-A chromatin assembly factor sufficient to form a functional de novo kinetochore [J ] . The Journal of Cell Biology , 2011 , 194 ( 2 ): 229 - 243 .

CAMAHORT R , LI B , FLORENS L , et al . Scm3 is essential to recruit the histone H3 variant Cse4 to centromeres and to maintain a functional kinetochore [J ] . Molecular Cell , 2007 , 26 ( 6 ): 853 - 865 .

DUNLEAVY E M , PIDOUX A L , MONET M , et al . A NASP (N1/N2)-related protein, Sim3, binds CENP-A and is required for its deposition at fission yeast centromeres [J ] . Molecular Cell , 2007 , 28 ( 6 ): 1029 - 1044 .

STOLER S , ROGERS K , WEITZE S , et al . Scm3, an essential Saccharomyces cerevisiae centromere protein required for G2/M progression and Cse4 localization [J ] . Proceedings of the National Academy of Sciences of the United States of America , 2007 , 104 ( 25 ): 10571 - 10576 .

WILLIAMS J S , HAYASHI T , YANAGIDA M , et al . Fission yeast Scm3 mediates stable assembly of Cnp1/CENP-A into centromeric chromatin [J ] . Molecular Cell , 2009 , 33 ( 3 ): 287 - 298 .

CHEN C C , DECHASSA M L , BETTINI E , et al . CAL1 is the Drosophila CENP-A assembly factor [J ] . The Journal of Cell Biology , 2014 , 204 ( 3 ): 313 - 329 .

LERMONTOVA I , KUHLMANN M , FRIEDEL S , et al . Arabidopsis KIN ETOCHORE NULL2 is an upstream component for centromeric histone H3 variant cenH3 deposition at centromeres [J ] . The Plant Cell , 2013 , 25 ( 9 ): 3389 - 3404 .

ZUO S , YADALA R , YANG F , et al . Recurrent plant-specific duplications of KNL2 and its conserved function as a kinetochore assembly factor [J ] . Molecular Biology and Evolution , 2022 , 39 ( 6 ): msac123 .

WOO H R , PONTES O , PIKAARD C S , et al . VIM1, a methylcytosine-binding protein required for centromeric heterochromatinization [J ] . Genes & Development , 2007 , 21 ( 3 ): 267 - 277 .

KRAFT E , BOSTICK M , JACOBSEN S E , et al . ORTH/VIM proteins that regulate DNA methylation are functional ubiquitin E3 ligases [J ] . The Plant Journal , 2008 , 56 ( 5 ): 704 - 715 .

ZÜRCHER J F , KLEEFELDT A A , FUNKE L F H , et al . Continuous synthesis of E . coli genome sections and Mb-scale human DNA assembly [J ] . Nature , 2023 , 619 ( 7970 ): 555 - 562 .

ZHONG L , ZHANG Q , LU N , et al . The conjugation-associated linear-BAC iterative assembling (CALBIA) method for cloning 2.1-Mb human chromosomal DNAs in bacteria [J ] . Cell Research , 2025 , 35 ( 4 ): 309 - 312 .

LAN T L , CHEN L G , WANG Y , et al . Genome synthesis in plants [J ] . Nature Reviews Bioengineering , 2025 : 326 .

SHAKIROV E V , CHEN J J L , SHIPPEN D E . Plant telomere biology: the green solution to the end-replication problem [J ] . The Plant Cell , 2022 , 34 ( 7 ): 2492 - 2504 .

MCKNIGHT T D , RIHA K , SHIPPEN D E . Telomeres, telomerase, and stability of the plant genome [J ] . Plant Molecular Biology , 2002 , 48 ( 4 ): 331 - 337 .

HOWELL S H . Molecular biology [M/OL ] . New York : Springer New York , 2014 [ 2025-09-01 ] . https://doi.org/10.1007/978-1-4614-7570-5 https://doi.org/10.1007/978-1-4614-7570-5 .

HU Y X , STILLMAN B . Origins of DNA replication in eukaryotes [J ] . Molecular Cell , 2023 , 83 ( 3 ): 352 - 372 .

STILLMAN B , DIFFLEY J F X , IWASA J H . Mechanisms for licensing origins of DNA replication in eukaryotic cells [J ] . Nature Structural & Molecular Biology , 2025 , 32 ( 7 ): 1143 - 1153 .

WHEELER E , BROOKS A M , CONCIA L , et al . Arabidopsis DNA replication initiates in intergenic, AT-rich open chromatin [J ] . Plant Physiology , 2020 , 183 ( 1 ): 206 - 220 .

FIORUCCI A S . AT the onset of DNA replication in Arabidopsis [J ] . Plant Physiology , 2020 , 183 ( 1 ): 19 - 20 .

JACOB Y , STROUD H , LEBLANC C , et al . Regulation of heterochromatic DNA replication by histone H3 lysine 27 methyltransferases [J ] . Nature , 2010 , 466 ( 7309 ): 987 - 991 .

COSTAS C , DE LA PAZ SANCHEZ M , STROUD H , et al . Genome-wide mapping of Arabidopsis thaliana origins of DNA replication and their associated epigenetic marks [J ] . Nature Structural & Molecular Biology , 2011 , 18 ( 3 ): 395 - 400 .

STROUD H , HALE C J , FENG S H , et al . DNA methyltransferases are required to induce heterochromatic re-replication in Arabidopsis [J ] . PLoS Genetics , 2012 , 8 ( 7 ): e1002808 .

EBERSOLE T A , ROSS A , CLARK E , et al . Mammalian artificial chromosome formation from circular alphoid input DNA does not require telomere repeats [J ] . Human Molecular Genetics , 2000 , 9 ( 11 ): 1623 - 1631 .

OKADA T , OHZEKI J I , NAKANO M , et al . CENP-B controls centromere formation depending on the chromatin context [J ] . Cell , 2007 , 131 ( 7 ): 1287 - 1300 .

NAKANO M , CARDINALE S , NOSKOV V N , et al . Inactivation of a human kinetochore by specific targeting of chromatin modifiers [J ] . Developmental Cell , 2008 , 14 ( 4 ): 507 - 522 .

WEUTS A , VOET T , VERBEECK J , et al . Telomere length homeostasis and telomere position effect on a linear human artificial chromosome are dictated by the genetic background [J ] . Nucleic Acids Research , 2012 , 40 ( 22 ): 11477 - 11489 .

KOUPRINA N , PETROV N , MOLINA O , et al . Human artificial chromosome with regulated centromere: a tool for genome and cancer studies [J ] . ACS Synthetic Biology , 2018 , 7 ( 9 ): 1974 - 1989 .

ZHU X T , LIU S C , YE T T , et al . Artificial chromosome reorganization reveals high plasticity of the budding and fission yeast genomes [J ] . Genome Biology , 2025 , 26 ( 1 ): 229 .

D’HALLUIN K , VANDERSTRAETEN C , VAN HULLE J , et al . Targeted molecular trait stacking in cotton through targeted double-strand break induction [J ] . Plant Biotechnology Journal , 2013 , 11 ( 8 ): 933 - 941 .

HOU L L , YAU Y Y , WEI J J , et al . An open-source system for in planta gene stacking by Bxb1 and Cre recombinases [J ] . Molecular Plant , 2014 , 7 ( 12 ): 1756 - 1765 .

KUMAR S , ALABED D , WORDEN A , et al . A modular gene targeting system for sequential transgene stacking in plants [J ] . Journal of Biotechnology , 2015 , 207 : 12 - 20 .

SRIVASTAVA V , THOMSON J . Gene stacking by recombinases [J ] . Plant Biotechnology Journal , 2016 , 14 ( 2 ): 471 - 482 .

ANAND A , WU E , LI Z , et al . High efficiency Agrobacterium -mediated site-specific gene integration in maize utilizing the FLP-FRT recombination system [J ] . Plant Biotechnology Journal , 2019 , 17 ( 8 ): 1636 - 1645 .

LI Y M , LI R Y , HAN Z G , et al . Recombinase-mediated gene stacking in cotton [J ] . Plant Physiology , 2022 , 188 ( 4 ): 1852 - 1865 .

ZONG Y , LIU Y J , XUE C X , et al . An engineered prime editor with enhanced editing efficiency in plants [J ] . Nature Biotechnology , 2022 , 40 ( 9 ): 1394 - 1402 .

LIU P , PANDA K , EDWARDS S A , et al . Transposase-assisted target-site integration for efficient plant genome engineering [J ] . Nature , 2024 , 631 ( 8021 ): 593 - 600 .

SUN C , LI H C , LIU Y J , et al . Iterative recombinase technologies for efficient and precise genome engineering across kilobase to megabase scales [J ] . Cell , 2025 , 188 ( 17 ): 4693 - 4710.e15 .

ELSHAMEY E A , YANG X M , YANG J Z , et al . Occurrence, biosynthesis, and health benefits of anthocyanins in rice and barley [J ] . International Journal of Molecular Sciences , 2025 , 26 ( 13 ): 6225 .

YE X D , AL-BABILI S , KLÖTI A , et al . Engineering the provitamin A (β-carotene) biosynthetic pathway into (carotenoid-free) rice endosperm [J ] . Science , 2000 , 287 ( 5451 ): 303 - 305 .

PAINE J A , SHIPTON C A , CHAGGAR S , et al . Improving the nutritional value of Golden Rice through increased pro-vitamin A content [J ] . Nature Biotechnology , 2005 , 23 ( 4 ): 482 - 487 .

LUDWIG M . Evolution of the C 4 photosynthetic pathway: events at the cellular and molecular levels [J ] . Photosynthesis Research , 2013 , 117 ( 1 ): 147 - 161 .

QIN K Z , YE X Y , LUO S S , et al . Engineering carbon assimilation in plants [J ] . Journal of Integrative Plant Biology , 2025 , 67 ( 4 ): 926 - 948 .

GUO K Y , YANG J , YU N , et al . Biological nitrogen fixation in cereal crops: Progress, strategies, and perspectives [J ] . Plant Communications , 2023 , 4 ( 2 ): 100499 .

LIU F , ZHAO Z H , FERNIE A R , et al . Towards establishing functional nitrogenase activities within plants [J/OL ] . Trends in Biotechnology , 2025 . ( 2025-05-28 )[ 2025-09-01 ] . https://doi.org/10.1016/j.tibtech.2025.04.020 https://doi.org/10.1016/j.tibtech.2025.04.020 .

WANG J , XIE Z X , MA Y , et al . Ring synthetic chromosome Ⅴ SCRaMbLE [J ] . Nature Communications , 2018 , 9 : 3783 .

CHOI S , NAH H J , CHOI S , et al . Heterologous expression of daptomycin biosynthetic gene cluster via Streptomyces artificial chromosome vector system [J ] . Journal of Microbiology and Biotechnology , 2019 , 29 ( 12 ): 1931 - 1937 .

TIAN T , WU X W , WU P P , et al . High-level expression of leghemoglobin in Kluyveromyces marxianus by remodeling the heme metabolism pathway [J ] . Frontiers in Bioengineering and Biotechnology , 2024 , 11 : 1329016 .

TRIPATHY S , DASSARMA B , BHATTACHARYA M , et al . Plant-based vaccine research development against viral diseases with emphasis on Ebola virus disease: a review study [J ] . Current Opinion in Pharmacology , 2021 , 60 : 261 - 267 .

JUGLER C , SUN H Y , NGUYEN K , et al . A novel plant-made monoclonal antibody enhances the synergetic potency of an antibody cocktail against the SARS-CoV-2 Omicron variant [J ] . Plant Biotechnology Journal , 2023 , 21 ( 3 ): 549 - 559 .