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1.

Wild emmer wheat, Triticum dicoccoides, the progenitor of modern tetraploid and hexaploid wheats, is an important resource for new variability for disease resistance genes. T. dicoccoides accession pau4656 showed resistance against prevailing leaf rust and stripe rust races in India and was used for developing stable introgression lines (IL) in T. durum cv Bijaga yellow and named as IL pau16068. F5 Recombinant inbred lines (F5 RILs) were developed by crossing IL pau16068 with T. durum cultivar PBW114 and RIL population was screened against highly virulent Pt and Pst pathotypes at the seedling and adult plant stages. Inheritance analyses revealed that population segregated for two genes for all stage resistance (ASR) against leaf rust, one ASR gene against stripe rust and three adult plant resistance (APR) genes for stripe rust resistance. For mapping these genes a set of 483 SSR marker was used for bulked segregant analysis. The markers showing diagnostic polymorphism in the resistant and susceptible bulks were amplified on all RILs. Single marker analysis placed all stage leaf rust resistance genes on chromosome 6A and 2A linked to the SSR markers Xwmc256 and Wpaus268, respectively. Likewise one all stage stripe rust resistance gene were mapped on long arm of chromosome 6A linked to markers 6AL-5833645 and 6AL-5824654 and two APR genes mapped on chromosomes 2A and 2B close to the SSR marker Wpaus268 and Xbarc70, respectively. The current study identified valuable leaf rust and stripe rust resistance genes effective against multiple rust races for deployment in the wheat breeding programme.

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2.
 The Yr15 gene of wheat confers resistance to the stripe rust pathogen Puccinia striiformis West., which is one of the most devastating diseases of wheat throughout the world. In the present study, molecular markers flanking the Yr15 gene of wheat have been identified using the near-isogenic-lines approach. RFLP screening of 76 probe-enzyme combinations revealed one polymorphic marker (Nor/TaqI) between the susceptible and the resistant lines. In addition, out of 340 RAPD primers tested, six produced polymorphic RAPD bands between the susceptible and the resistant lines. The genetic linkage of the polymorphic markers was tested on segregating F2 population (123 plants) derived from crosses between stripe rust-susceptible Triticum durum wheat, cv D447, and a BC3F9 resistant line carrying Yr15 in a D447 background. A 2.8-kb fragment produced by the Nor RFLP probe and a 1420-bp PCR product generated by the RAPD primer OPB13 showed linkage, in coupling, with the Yr15 gene. Employing the standard maximum-likelihood technique it was found that the order OPB13 1420 Yr15Nor1 on chromosome 1B appeared to be no less than 1000-times more probable than the closest alternative. The map distances between OPB13 1420 Yr15Nor1 are 27.1 cM and 11.0 cM for the first and second intervals, respectively. The application of marker-assisted selection for the breeding of new wheat cultivars with the stripe rust resistance gene is discussed. Received: 27 February 1997/Accepted: 7 March 1997  相似文献   

3.

Key Message

Rapid LD decay in wild emmer population from Israel allows high-resolution association mapping. Known and putative new stripe rust resistance genes were found.

Abstract

Genome-wide association mapping (GWAM) is becoming an important tool for the discovery and mapping of loci underlying trait variation in crops, but in the wild relatives of crops the use of GWAM has been limited. Critical factors for the use of GWAM are the levels of linkage disequilibrium (LD) and genetic diversity in mapped populations, particularly in those of self-pollinating species. Here, we report LD estimation in a population of 128 accessions of self-pollinating wild emmer, Triticum turgidum ssp. dicoccoides, the progenitor of cultivated wheat, collected in Israel. LD decayed fast along wild emmer chromosomes and reached the background level within 1 cM. We employed GWAM for the discovery and mapping of genes for resistance to three isolates of Puccinia striiformis, the causative agent of wheat stripe rust. The wild emmer population was genotyped with the wheat iSelect assay including 8643 gene-associated SNP markers (wheat 9K Infinium) of which 2,278 were polymorphic. The significance of association between stripe rust resistance and each of the polymorphic SNP was tested using mixed linear model implemented in EMMA software. The model produced satisfactory results and uncovered four significant associations on chromosome arms 1BS, 1BL and 3AL. The locus on 1BS was located in a region known to contain stripe rust resistance genes. These results show that GWAM is an effective strategy for gene discovery and mapping in wild emmer that will accelerate the utilization of this genetic resource in wheat breeding.  相似文献   

4.
Stripe rust (Puccinia striiformis f. sp. tritici) is one of the major devastating disease which causes large reduction in wheat yield. T. monococcum is an attractive diploid species for gene discovery in wheat with smaller genome size of 5700 Mb compared to 17,300 Mb of bread wheat. An adult plant stripe rust resistance QTL QYrtm.pau-2A was mapped on chromosome 2A flanked by two SSR markers Xwmc170 and Xwmc407. In the present study, two gene based markers Pau_Ta2AL_Gene45 and Pau_Ta2AL_Gene54 developed from 2A specific ESTs were found to map close to QYrtmpau-2A to narrow down the region for candidate gene identification. Utilizing sequence information of these two markers, four BAC clones were identified from the Minimum Tiling Path of 2AL assembly and were sequenced. SSR markers were designed from these BAC sequences and mapped to chromosome 2A. A 50 Mb region of wheat chromomse 2A was identified to harbor stripe rust resistance gene of T. monococcum. Gene based markers identified in the present investigation can be used for marker assisted introgression of QYrtm.pau-2A from T. monococcum to cultivated wheat.  相似文献   

5.
D Bai  D R Knott 《Génome》1994,37(3):405-409
Six accessions of Triticum turgidum var. dicoccoides L. (4x, AABB) of diverse origin were tested with 10 races of leaf rust (Puccinia recondita f.sp. tritici Rob. ex Desm.) and 10 races of stem rust (P. graminis f.sp. tritici Eriks. &Henn.). Their infection type patterns were all different from those of lines carrying the Lr or Sr genes on the A or B genome chromosomes with the same races. The unique reaction patterns are probably controlled by genes for leaf rust or stem rust resistance that have not been previously identified. The six dicoccoides accessions were crossed with leaf rust susceptible RL6089 durum wheat and stem rust susceptible 'Kubanka' durum wheat to determine the inheritance of resistance. They were also crossed in diallel to see whether they carried common genes. Seedlings of F1, F2, and BC1F2 generations from the crosses of the dicoccoides accessions with RL6089 were tested with leaf rust race 15 and those from the crosses with 'Kubanka' were tested with stem rust race 15B-1. The F2 populations from the diallel crosses were tested with both races. The data from the crosses with the susceptible durum wheats showed that resistance to leaf rust race 15 and stem rust race 15B-1 in each of the six dicoccoides accessions is conferred by a single dominant or partially dominant gene. In the diallel crosses, the dominance of resistance appeared to be affected by different genetic backgrounds. With one exception, the accessions carry different resistance genes: CI7181 and PI 197483 carry a common gene for resistance to leaf rust race 15. Thus, wild emmer wheat has considerable genetic diversity for rust resistance and is a promising source of new rust resistance genes for cultivated wheats.  相似文献   

6.
抗条锈病基因Yr69对我国小麦条锈菌(Puccinia striiformis f. sp. tritici)小种具有广谱抗性,在小麦抗条锈病育种中具有重要价值。为提高分子标记辅助选择育种的效率,加快Yr69在小麦抗病育种中的应用,本研究利用条锈菌小种CYR34对包含340个小麦家系的‘Taichung29/CH7086’F9代RIL(Recombinant inbred line)群体进行接种鉴定,并利用BSA-SNP(Bulked segregant analysis-single nucleotide polymorphism)技术对其抗条锈病基因进行了重新定位。抗病鉴定结果显示,RIL群体中抗感病家系的数量呈双峰分布,‘CH7086’的条锈病抗性受一个主效位点控制。BSA-SNP基因分型结果表明,多态性SNP主要集中于小麦2AS染色体末端0~30Mb的染色体区段。在该基因组区段开发了208个SSR分子标记,利用抗感病小群体从中筛选到14个与Yr69连锁的分子标记。利用14个标记对340个RIL家系进行PCR扩增和分子作图,将Yr69定位于2AS111和2AS171之间约7.76...  相似文献   

7.
Several new races of the stripe rust pathogen have become frequent throughout the wheat growing regions of the United States since 2000. These new races are virulent to most of the wheat seedling resistance genes limiting the resistance sources that can be used to combat this pathogen. High-temperature adult-plant (HTAP) stripe rust resistance has proven to be more durable than seedling resistance due to its non-race-specific nature, but its use is limited by the lack of mapping information. We report here the identification of a new HTAP resistance gene from Triticum turgidum ssp. dicoccoides (DIC) designated as Yr36. Lines carrying this gene were susceptible to almost all the stripe rust pathogen races tested at the seedling stage but showed adult-plant resistance to the prevalent races in California when tested at high diurnal temperatures. Isogenic lines for this gene were developed by six backcross generations. Field tests in two locations showed increased levels of field resistance to stripe rust and increased yields in isogenic lines carrying the Yr36 gene compared to those without the gene. Recombinant substitution lines of chromosome 6B from DIC in the isogenic background of durum cv. Langdon were used to map the Yr36 gene on the short arm of chromosome 6B completely linked to Xbarc101, and within a 2-cM interval defined by PCR-based markers Xucw71 and Xbarc136. Flanking locus Xucw71 is also closely linked to the grain protein content locus Gpc-B1 (0.3-cM). Marker-assisted selection strategies are presented to improve stripe rust resistance and simultaneously select for high or low Gpc-B1 alleles.  相似文献   

8.
Quantitative trait loci of stripe rust resistance in wheat   总被引:1,自引:0,他引:1  

Key message

Over 140 QTLs for resistance to stripe rust in wheat have been published and through mapping flanking markers on consensus maps, 49 chromosomal regions are identified.

Abstract

Over thirty publications during the last 10 years have identified more than 140 QTLs for stripe rust resistance in wheat. It is likely that many of these QTLs are identical genes that have been spread through plant breeding into diverse backgrounds through phenotypic selection under stripe rust epidemics. Allelism testing can be used to differentiate genes in similar locations but in different genetic backgrounds; however, this is problematic for QTL studies where multiple loci segregate from any one parent. This review utilizes consensus maps to illustrate important genomic regions that have had effects against stripe rust in wheat, and although this methodology cannot distinguish alleles from closely linked genes, it does highlight the extent of genetic diversity for this trait and identifies the most valuable loci and the parents possessing them for utilization in breeding programs. With the advent of cheaper, high throughput genotyping technologies, it is envisioned that there will be many more publications in the near future describing ever more QTLs. This review sets the scene for the coming influx of data and will quickly enable researchers to identify new loci in their given populations.  相似文献   

9.
10.
Stripe rust resistance was identified in Triticum vavilovii (T. vavilovii Aus22498)-derived Russian wheat aphid (RWA)-resistant germplasm. Inheritance studies indicated monogenic control of resistance. The resistance gene was tentatively designated as Yrvav and was located on chromosome 1B by monosomic analysis. A close association (1.5±0.9% recombination) of Yrvav with a T. vavilovii-derived gliadin allele (Gli-B1vav) placed it in chromosome arm 1BS. Yrvav was allelic with Yr10. Tests with Yr10 avirulent and virulent pathotypes showed that Yrvav and Yr10 possess identical pathogenic specificity. Yrvav and Yr10 showed close genetic associations with alternate alleles at the Xpsp3000 (microsatellite marker), Gli-B1 and Rg1 loci. Based on these observations Yrvav was named as Yr10vav. The close association between Xpsp3000 and Gli-B1 was also confirmed. The Yr10vav-linked Xpsp3000 allele (285 bp) was not present in 65 Australian cultivars, whereas seven Australian wheats lacking Yr10 carried the same Xpsp3000 allele (260 bp) as Yr10 carrying wheat cultivar Moro. Xpsp3000 and/or Gli-B1 could be used in marker-assisted selection for pyramiding Yr10vav or Yr10 with other stripe rust resistance genes. Yr10vav was inherited independently of the T. vavilovii-derived RWA resistance. Received: 5 December 2000 / Accepted: 3 April 2001  相似文献   

11.
Growing resistant wheat varieties is a key method of controlling two important wheat diseases, leaf rust and stripe rust. We analyzed quantitative trait loci (QTL) to investigate adult plant resistance (APR) to these rusts, using 141 F5 RILs derived from the cross ‘Avocet-YrA/Francolin#1’. Phenotyping of leaf rust resistance was conducted during two seasons at Ciudad Obregon, Mexico, whereas stripe rust was evaluated for two seasons in Toluca, Mexico, and one season in Chengdu, China. The genetic map was constructed with 581 markers, including diversity arrays technology and simple sequence repeat. Significant loci for reducing leaf rust severity were designated QLr.cim-1BL, QLr.cim-3BS.1, QLr.cim-3DC, and QLr.cim-7DS. The six QTL that reduced stripe rust severity were designated QYr.cim-1BL, QYr.cim-2BS, QYr.cim-2DS, QYr.cim-3BS.2, QYr.cim-5AL, and QYr.cim-6AL. All loci were conferred by Francolin#1, with the exception of QYr.cim-2DS, QYr.cim-5AL, and QYr.cim-6AL, which were derived from Avocet-YrA. Closely linked markers indicated that the 1BL locus was the pleiotropic APR gene Lr46/Yr29. QYr.cim-2BS was a seedling resistance gene designated as YrF that conferred intermediate seedling reactions and moderate resistance at the adult plant stage in both Mexican and Chinese environments. Significant additive interactions were detected between the six QTL for stripe rust, but not between the four QTL for leaf rust. Furthermore, we detected two new APR loci for leaf rust in common wheat: QLr.cim-3BS.1 and QLr.cim-7DS.  相似文献   

12.
Yang WY  Yu Y  Zhang Y  Hu XR  Wang Y  Zhou YC  Lu BR 《Hereditas》2003,139(1):49-55
Stripe rust is one of the most destructive diseases for wheat crops in China. Two stripe rust physiological strains, i.e. CYR30 (intern. name: 175E191) and CYR31 (intern. name: 293E175) have been the dominant and epidemic physiological strains since 1994. One Aegilops tauschii accession (SQ-214) from CIMMYT was found immune from or highly resistant to Chinese new stripe rust races CYR30 and CYR31 at adult stage. SQ-214 was crossed with a highly susceptible Ae. tauschii accession As-80. Analysis of data from F1-F2 populations of SQ-214/As-80 revealed that the resistance was controlled by a single dominant gene. To exploit the resistance for wheat breeding, SQ-214 was crossed with Chinese Spring (CS) and backcrossed by two Chinese commercial wheat varieties MY26 and SW3243. The resistance from SQ-214 was suppressed in the F1 hybrids (CS/SQ-214) and the F2 population of CS/SQ-214//MY26. However, the resistance of SQ-214 was expressed in several F2 individuals of CS/SQ-214//SW3243. Eleven advanced lines with high level of resistance to the Chinese stripe rust CYR30 and CYR31 have been developed. This result suggested that SW3243 does not suppress the expression of the Chinese stripe rust and should be used as wheat germplasm for exploiting resistance of Ae. tauschii in wheat breeding. The gliadin electrophoretic pattern of the eleven advanced lines with high stripe rust resistances was compared with their parents SQ-214, CS and SW3243 by acid polyacrylamide gel electrophoresis. The omega-gliadin bands of Gli-Dt1 in Ae. tauschii SQ-214 were transferred to some advanced lines and freely expressed in common wheat genetic background. One of advanced lines possesses a null Gli-D1 allele, where the omega-gliadin bands encoding by the Gli-D1 allele were absent. The potential utilization of this advanced line for wheat quality and stripe rust resistance breeding is also discussed in this paper.  相似文献   

13.
Stripe rust resistance in the German winter wheat cv. Alcedo has been described as durable, the resistance having remained effective when grown extensively in Germany and Eastern Europe between 1975 and 1989. Genetic characterisation of field resistance in a cross between Alcedo and the stripe rust susceptible UK winter wheat cv. Brigadier identified two major QTL in Alcedo located on the long arms of chromosomes 2D (QPst.jic-2D) and 4B (QPst.jic-4B). Stripe rust resistance was evaluated by measuring the extent of fungal growth, percentage infection (Pi) and the necrotic/chlorotic response of the plant to infection, infection type (IT). Both QPst.jic-2D and QPst.jic-4B contributed significantly to the reduction in stripe rust infection (Pi), with QPst.jic-2D explaining up to 36.20% and QPst.jic-4B 28.90% of the phenotypic variation measured for Pi. Both QTL were identified by the IT phenotypic scores, with QPst.jic-2D in particular being associated with a strong necrotic phenotype (low IT), QPst.jic-2D explaining up to 53.10% of IT phenotypic variation and QPst.jic-4B 22.30%. In addition, two small effect QTL for field stripe rust resistance were identified in Brigadier, QPst.jic-1B on the long arm of chromosome 1B and QPst.jic-5A on the short arm of chromosome 5A. The influence of QPst.jic-1B was primarily seen with the Pi phenotype, contributing up to 13.10% of the explained phenotypic variation. QPst.jic-5A was only detected using an approximate multiple-QTL model and selecting markers linked to the major effect QTL, QPst.jic-2D and QPst.jic-4B as co-factors. Seedling stripe rust resistance was also mapped in the cross, which confirmed the location of Yr17 from Brigadier to the short arm of chromosome 2A. A seedling expressed QTL was also located in Alcedo that mapped to the same location as the field stripe rust resistance QPst.jic-2D.  相似文献   

14.
Following the appearance of stripe rust in South Africa in 1996, efforts have been made to identify new sources of durable resistance. The French cultivar Cappelle-Desprez has long been considered a source of durable, adult plant resistance (APR) to stripe rust. As Cappelle-Desprez contains the seedling resistance genes Yr3a and Yr4a, wheat lines were developed from which Yr3a and Yr4a had been removed, while selecting for Cappelle-Desprez derived APR effective against South African pathotypes of the stripe rust fungus, Puccinia striiformis f. sp. tritici. Line Yr16DH70, adapted to South African wheat growing conditions, was selected and crossed to the stripe rust susceptible cultivar Palmiet to develop a segregating recombinant inbred line mapping population. A major effect QTL, QYr.ufs-2A was identified on the short arm of chromosome 2A derived from Cappelle-Desprez, along with three QTL of smaller effect, QYr.ufs-2D, QYr.ufs-5B and QYr.ufs-6D. QYr.ufs-2D was located within a region on the short arm of chromosome 2D believed to be the location of the stripe rust resistance gene Yr16. An additional minor effect QTL, QYr.ufs-4B, was identified in the cv. Palmiet. An examination of individual RILs carrying single or combinations of each QTL indicated significant resistance effects when QYr.ufs-2A was combined with the three minor QTL from Cappelle-Desprez, and between QYr.ufs-2D and QYr.ufs-5B.  相似文献   

15.
ABSTRACT: BACKGROUND: Non-host resistance (NHR) confers plant species immunity against the majority of microbialpathogens and represents the most robust and durable form of plant resistance in nature. Asone of the main genera of rust fungi with economic and biological importance, Puccinia infects almost all cereals but is unable to cause diseases on legumes. Little is known about themechanism of this kind of effective defense in legumes to these non-host pathogens. RESULTS: In this study, the basis of NHR in broad bean (Vicia faba L.) against the wheat stripe rustpathogen, Puccinia striiformis f. sp. tritici (Pst), was characterized. No visible symptomswere observed on broad bean leaves inoculated with Pst. Microscopic observations showedthat successful location of stomata and haustoria formation were significantly reduced in Pstinfection of broad bean. Attempted infection induced the formation of papillae, cell wallthickening, production of reactive oxygen species, callose deposition and accumulation ofphenolic compounds in plant cell walls. The few Pst haustoria that did form in broad beancells were encased in reactive oxygen and callose materials and those cells elicited cell death.Furthermore, a total of seven defense-related genes were identified and found to be upregulatedduring the Pst infection. CONCLUSIONS: The results indicate that NHR in broad bean against Pst results from a continuum of layereddefenses, including basic incompatibility, structural and chemical strengthening of cell wall,posthaustorial hypersensitive response and induction of several defense-related genes,demonstrating the multi-layered feature of NHR. This work also provides useful informationfor further determination of resistance mechanisms in broad bean to rust fungi, especially theadapted important broad bean rust pathogen, Uromyces viciae-fabae, because of strongsimilarities and association between NHR of plants to unadapted pathogens and basalresistance of plants to adapted pathogens.  相似文献   

16.
Stripe rust caused by the fungus Puccinia striiformis f. sp. tritici (Pst) may decrease wheat yield significantly in severe outbreaks. The most cost-effective and environmentally friendly approach to reduce yield losses due to rust diseases is deployment of effective resistant genes in wheat cultivars. The causal agents evolve and may break existing resistant sources as well. Therefore, long-term conventional breeding strategies and the ongoing evolution of pathogen populations in the region would put the success of breeding programmes at risk so that there is always a need for speeding up the process of germplasm enhancement through production of doubled-haploid breeding materials. In this study, we aimed at introgression of stripe rust resistance trait from three genotypes (Flanders, Martonvasar-17 (MV17) and Bersee) into a widely adapted cultivar “Ghods”. Positively selected F2BC2 progenies of three backcrossing schemas, i.e. (i) Flanders/3*Ghods; (ii) Ghods*3/MV17; and (iii) Hybride-de-bersee/3*Ghods, were used to produce three small-size doubled-haploid populations via wheat × Maize pollination methodology. The doubled-haploid populations were examined against two predominantly isolates of P. striiformis f. sp. tritici (Pst) i.e. 6E134A+ and 6E2A+Yr27+ and the screening revealed that 44 and 52 of the progenies are resistant to the above-mentioned isolates, respectively. Field data have shown that the stripe rust resistance doubled-haploid germplasm are comparable to local check cultivars in yield and earliness.  相似文献   

17.
18.
西科麦2028是地理远缘小麦材料的杂交后代,具有突出的抗条锈病性能。为了解西科麦2028对小麦条锈病的抗性遗传规律,以西科麦2028和铭贤169的杂交群体为研究对象,采用我国目前小麦条锈菌流行小种CYR31、CYR32、CYR33、Su11-4对供试群体进行成株期接种,分析杂交后代的抗病性及分布情况。结果表明:西科麦2028对CYR31的抗病性由3对显性基因控制;对CYR32由2对显性和1对隐性基因控制;对CYR33由1对显性基因控制;对Su11-4由1对显性和1对隐性基因控制。  相似文献   

19.
Wollaroi, an Australian durum wheat cultivar, produced a low stripe rust response and the alternative parent Bansi was highly susceptible. The Wollaroi/Bansi recombinant inbred line (RIL) population was phenotyped across three consecutive crop seasons. A genetic map of the Wollaroi/Bansi RIL population comprising 799 markers (diversity arrays technology and simple sequence repeat markers) was used to determine the genomic location of stripe rust resistance genes carried by the cultivar Wollaroi. Composite interval mapping detected three consistent quantitative trait loci (QTL) in chromosomes 2A, 3B and 5B. These QTL were named QYr.sun-2A, QYr.sun-3B and QYr.sun-5B. Another QTL, QYr.sun-1B, was detected only in the 2009 crop season. QTL in chromosomes 1B, 2A, 3B and 5B explained on average 6, 9.3, 26.7 and 8.7 %, respectively, of the variation in stripe rust response. All QTL were contributed by Wollaroi. RILs carrying these QTL singly produced intermediate stripe rust severities ranging from 46.2 to 55.7 %, whereas RILs with all four QTL produced the lowest disease severity (34.3 %). The consistently low stripe rust response of Wollaroi for 20 years demonstrated the durability of the resistance loci involved. The QTL combination detected in this study is being transferred to common wheat.  相似文献   

20.
Dose–response relationship between resistance of wheat seedlings (Triticum aestivum, cultivar Zarya) to Erysiphe graminis f. sp. tritici Marchal. (Syn. Blumeria graminis), a causal organism of wheat powdery mildew and exogenous zeatin has been investigated. Two-week-old seedlings were inoculated with the pathogen. Zeatin or zeatinriboside were added to the nutrient solution immediately after inoculation. The dose–response curve of cytokinin in the most cases was multiphasic, with peaks of increased susceptibility occurring at 0.25–1.5 and 1.5–9 μM cytokinin, separated by a region of increased resistance at 0.5–3 μM cytokinin. The change in mineral nutrition or simultaneous treatment with thidiazuron revealed alterations of the dose–response curve ranging from a curve with maximum of resistance to a curve with maximum of susceptibility. Both multiphase nature of dose–response and its variability were proposed as possible explanations for earlier observed discrepancies in experimental data on modification of disease resistance by cytokinins. A mathematical model for two metabolic processes with substrate inhibition connected in-series was suggested to explain the multiphase dose–response. In this model, the product of the first reaction was used as substrate for the second reaction. Numerical experiments showed the changes in the shape of dose–response curve with changes in parameters dependent of cytokinin metabolism.  相似文献   

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