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Early spontaneous diploidization of maternal maize haploids generated by in vivo haploid induction

Early spontaneous diploidization of maternal maize haploids generated by in vivo haploid induction
Early spontaneous diploidization of maternal maize haploids generated by in vivo haploid induction

Early spontaneous diploidization of maternal maize haploids generated by in vivo haploid induction

Penghao Wu ?Jiaojiao Ren ?Liang Li ?

Shaojiang Chen

Received:23December 2013/Accepted:27May 2014/Published online:11June 2014óSpringer Science+Business Media Dordrecht 2014

Abstract The production of doubled haploid (DH)lines has become a key technology in maize (Zea mays L.)research and breeding.However,most of the haploid plants are sterile and in many cases arti?cial chromosome doubling involves the use of costly and toxic chemicals.Here,we report a special kind of doubled haploid named the early doubled haploid (EH)that was generated directly by in vivo haploid induc-tion.We found 83EH plants induced from the hybrid Zhengdan958,55families of its F 2:3population and the parental lines,all of which were con?rmed to be homozygous diploids via ?ow cytometry and 104SSR markers.The progeny of EH 0(EH 1)behaved in the same manner and showed the same potentialities as the parents of Zheng58and Chang7-2.EH plants were also detected in other genetic backgrounds at a frequency of 1–3.5%based on the total number of haploid plants.Because the EH lines exhibited completely fertility and

were obtained from induction directly in one step,they could be used in DH breeding as a new breeding strategy.According to our observations,it is likely that spontaneous doubling in EH occurred during embryo development when haploid induction.The possible mechanism of EH is also discussed.

Keywords Maize áChromosome doubling áEarly doubled haploids áHaploid diploidization áHaploid induction

Abbreviations DH Doubled haploids EH Early doubled haploids KOC Kernel oil content

SSR Simple sequence repeat

Introduction

The development of homozygous inbred lines is important in the breeding of maize and many other crops.Breeders and researchers have traditionally produced inbred lines by sel?ng heterozygous mate-rials for ?ve to six generations,but this approach is expensive and time consuming (Eder and Chalyk 2002).Breeding with haploids can result in the creation of pure breeding lines (Szarejko and Forster 2007;Chang and Coe 2009;Geiger and Gordillo 2009)in fewer seasons.

Electronic supplementary material The online version of this article (doi:10.1007/s10681-014-1166-5)contains supple-mentary material,which is available to authorized users.Penghao Wu and Jiaojiao Ren contributed equally to this work.P.Wu áJ.Ren áL.Li áS.Chen (&)

National Maize Improvement Center of China,China Agricultural University,No.2Yuanmingyuan West Road,Haidian District,Beijing 100193,China e-mail:shaoj@https://www.doczj.com/doc/5c14933961.html, P.Wu

e-mail:wupenghao@https://www.doczj.com/doc/5c14933961.html,

Euphytica (2014)200:127–138DOI 10.1007/s10681-014-1166-5

Haploid induction and identi?cation are not prob-lematic with the selection of ef?cient haploid induc-ers,such as WS14(Lashermes and Beckert1988), ZMS(Chalyk1994),MHI(Eder and Chalyk2002), CAUHOI(Liu and Song2000)and RWS(Ro¨ber et al. 2005).Haploids produced by these inducers are completely male-serile except for sectors in which doubling of chromosomes has taken place.There are two traditional methods for obtaining fertile haploids. One is spontaneous haploid doubling,for which doubling rates ranging from0to21.4%have been reported in various maize genotypes(Chase1952; Chalyk1994;Barnabas et al.1999).The other is arti?cial chromosome doubling,which can duplicate the chromosome set of haploid plants effectively using doubling agents or physical stress during treatment. Many studies have focused on the improvement of arti?cial doubling ef?ciency(Stadler et al.1989;Wan et al.1991;Ha¨ntzschel and Weber2010).Colchicine is the most widely applied arti?cial chromosome doubling agent(Gayen et al.1994;Deimling et al. 1997).Despite the effectiveness of colchicine in arti?cial chromosome doubling and its suitability for large-scale DH line production(Gayen et al.1994; Barnabas et al.1999),in many cases its high cost, toxicity and labor intensiveness present major chal-lenges for DH line production.In addition,relatively few seeds are produced on each haploid and they therefore need to be propagated for another generation before they can be used in breeding applications.Also, as many of the induced haploids have a certain spontaneous doubling ability both in male and female, arti?cial chromosome duplication is not necessary for DH line production from germplasm with high proportion of fertile haploids(Kleiber et al.2012).

Several researchers have reported the partial spon-taneous duplication of haploids(Zabirova et al.1993; Chalyk1994;Geiger et al.2006;Geiger and Scho¨nle-ben2011).They found variations in partial fertility recovery rates in different materials.Regarding the mechanism of spontaneous duplication of haploid plants,Testillano et al.(2004)studied chromosome doubling in early microspore embryogenesis in vitro and determined that fusion of nuclei likely leads to chromosome doubling.Moreover,spontaneous hap-loid doubling is always considered independently with haploid induction when taking into account double-fertilization mechanisms.Several mutants were found in efforts to describe the exact double fertilization process,like feronia(Huck et al.2003),sirene (Rotman et al.2003),ig1(indeterminate gameto-phyte1)(Evans2007)and cdka(Nowack et al.2006), and haploid inducers can also be considered as original mutant lines to help in the analysis of double-fertilization mechanisms(Li et al.2009).The rela-tionship between double-fertilization and spontaneous doubling is unclear.

In exception to this,the frequency of spontaneous chromosome doubling in some elite cultivars may be high enough to skip the doubling step and use the directly-obtained DHs(Kasha and Kao1970;Hoek-stra et al.1993).In earlier study,researchers found that some haploids could spontaneously double in early development(Chalyk1994).However,this phenom-enon was not systematically studied and people did not pay much attention to its value in breeding program for a long time.In this study,these early doubled haploids were named as‘EH’and could be obtained in one step for use in maize breeding as pure lines.The objectives of this study were:(1)to report the phenomenon of early diploidization during in vivo haploid induction in maize;(2)to examine the homozygosity of the EH lines;(3)to provide insight into the biological mechanisms of spontaneous haploid doubling. Materials and methods

Haploid induction

CAUHOI(Liu and Song2000)was used as the haploid inducer line.The maize single-cross hybrid Zheng-dan958,55families of its F2:3population and the parental lines Zheng58and Chang7-2were used as the female parents.Zhengdan958was developed by the Henan Academy of Agriculture Science.The crosses were made manually at the Shangzhuang Experimen-tal Station in Beijing in July2010.The harvested putative haploids were screened by the marker gene R1-nj(Nanda and Chase1966)and planted in Beijing in summer of2011.All male-fertile haploid plants and haploid plants with tassel sectors were selfed.

Plant classi?cation and morphological observation

The inducer line CAUHOI is homozygous for the dominant marker gene plant purple coloration gene (Liu and Song2000),which leads to purple stalks and

leaves.Normally,the F1plants are vigorous because of heterosis and have purple stalks and leaves,whereas the haploid plants are shorter,with slender weak stems and narrower leaves,and grow slowly.However,a new kind of plant was observed before the?owering stage,showing green stalks and leaves but no heter-osis,which was obviously not from contamination during the haploid induction process.We supposed these plants were doubled haploids that were doubled at an early stage.Hence,we named them early doubled haploids(EH).To study this new kind of plant,the plants from the putative haploids were classi?ed into three groups:(1)haploid plants,which were short with green stalks and upright green leaves;(2)hybrid plants,which were strong with purple stalks and leaves;(3)EH plants,which had green stalks and leaves but were much stronger than haploid plants. During the?owering stage,all plants with completely fertile or partially fertile tassels were selfed.Morpho-logical traits,such as plant height,ear height,tassel length and tassel branch number were measured at ?owering time.Ear length,ear width and the number of selfed kernels were measured after harvest.To distinguish between EH and other DH lines,the DH lines generated from fertile haploid plants are named normal doubled haploids(DH)hereafter.The fertile EH plants were designated EH0and the harvested kernels were designated EH1.The fertile haploid plants were designated DH0and the harvested kernels were designated DH1.

Ploidy level determination using?ow cytometry

To complement visual scoring of ploidy levels,three categories of plants at?owering time were examined with?ag leaves by?ow cytometry as described by Palomino et al.(2008)using a CAII?ow cytometer (Partec GmbH,Mu¨nster,Germany):(i)10normal diploids and10completely male-sterile haploids as a control,(ii)137DH0plants,and(iii)83EH0plants. The DPAC software was used for data analysis. Nuclear ploidy was represented by C values,with1C representing haploid,2C representing diploid,and so on(Ha¨ntzschel and Weber2010).

Simple sequence repeat(SSR)marker analysis

A total of83EH0plants and the parental lines of Zheng58and Chang7-2were chosen for SSR analysis.Maize genomic DNA was extracted from ?ag leaves at?owering time using the CTA

B proce-dure(Saghai-Maroof et al.1984).A total of367SSR primer sequences were screened for polymorphisms between Zheng58and Chang7-2,Zheng58and CAU-HOI,and Chang7-2and CAUHOI,respectively.Of these SSR markers,104SSR primer sequences showing clear polymorphisms in Zheng58and Chang7-2,107in Zheng58and CAUHOI,and105in Chang7-2and CAUHOI(covering all10chromo-somes)were used for SSR primer sequences analysis. All SSR primer sequences were obtained from the MaizeGDB database(https://www.doczj.com/doc/5c14933961.html,/ssr. php).DNA ampli?cation and polymorphism identi?-cation were performed as described by Ninamango-Ca′rdenas et al.(2003).

Evaluation of EH1lines for agronomic traits

Self-pollinated ears(EH1)from EH0plants were manually harvested and ear traits were recorded on an individual plant basis.The ears were then grown ear-to-row in an observation nursery with up to12plants per row to visually evaluate plant uniformity.A set of 33EH1lines from the Zhengdan958haploids and the two parental lines were evaluated under normal conditions during the summer of2012in Beijing.A completely randomized block design was used,with three replications for each genotype.In each block, plants were sown in single-rows,2.5-m long,with a density of60,000plants/ha.Uni?ed management measures,such as irrigation,fertilization and weed cutting were applied during the whole growth period. Data were recorded on the following traits:plant height,ear height,tassel length,tassel branch number, ear width and ear length.Nine plants in the middle of each row were chosen for data collection. Statistical analysis

For the agronomic traits(plant height,ear height, tassel length,tassel branch number,kernel number,ear width and ear length),analysis of variance(ANOVA) was performed by a generalized linear model(GLM) in SPSS(SPSS11.5for Windows)software.Means and variances of each EH and the controls were calculated.Mean comparison between EH and the checks was performed using Scheffe’s test at5% level of signi?cance.To con?rm the homogeneity of

the EH1lines,the intra-variances(average of within-line variance)of the EH1lines and controls were obtained and evaluated using F-statistics.The parents, Zheng58and Chang7-2,served as controls.

EH0in other inbred lines

To evaluate the EH frequency in other different materials,seven other elite inbred lines were also crossed with CAUHOI in the winter of2011in Hainan,including4F1,B73,By815,Dan598,8701, Qi319and Xu178.The putative haploids of these materials(including Zheng58and Chang7-2)were planted in Beijing and Gansu in the summer of2012. Other haploids from Zhengdan958were treated with colchicine and also planted in Beijing in2012,the methods of treatment were refer to Ha¨ntzschel and Weber(2010).The EH0frequency was recorded for these materials and all fertile plants were selfed.EH0 plants were also con?rmed by?ow cytometry and SSR markers.The proportion of EH0plants among haploid plants was called as EDR(early doubled rate)and the proportion of EH0plants among the seeds produced by the cross was called as EHI(early doubled haploid induction rate).A set of5EH1lines from each inbred line(Zheng58,Chang7-2,B73and8701)including the parents and their DH1lines were evaluated under normal conditions using a randomized complete block design(RCBD)with3replications in the winter of 2012in Hainan(other?ve inbred lines were not be evaluated because of lack of DH lines).Each line was grown in single-rows,2.5-m long,with a density of 60,000plants/ha.Uni?ed management measures were applied during the whole growth period.Data were recorded on the following traits:plant height,ear height,tassel length and tassel branch number.Nine plants in the middle of each row were chosen for data collection.The methods of statistical analysis were the same with way above.

Results

Morphology of EH0

The performances of EH0plants were clearly different from either the normal haploids or hybrid plants.First, the EH0plants were more vigorous than normal haploids.The average values for plant height and ear height in EH0plants were188and77cm,which were signi?cantly higher than the haploids(Fig.1a; Table1).Second,no heterosis was detected in the EH0plants but it was obvious in the hybrid plants, which had averages of247cm for plant height and 144cm for ear height(Fig.1a).In addition,hybrid plants had purple leaves and stalks as they were induced by CAUHOI with the ABP1R marker genes (A1:anthocyanin gene;B:plant color strengthening gene;P1:plant purple coloration gene;R:plant aleurone color gene)(Chase1969;Liu and Song 2000),but no purple EH0plants were detected (Fig.1a).Moreover,most DH0plants displayed only low levels of female and male fertility whereas all the EH0plants showed normal fertility.The means of ear width and length in EH0ears from Zhengdan958were about2–3times greater than in DH0ears(Fig.1b; Table1).The seed setting was much better in EH0ears than in DH0ears.The EH0ear had averagely more than100kernels,whereas DH0ear had averagely less than10kernels(Fig.1b;Table1;Supplementary Table1).

Ploidy level of EH0plants

Haploid plants had a signi?cant peak at the C position (Fig.2a),while normal diploid plants of the inbred line Zheng58had a signi?cant peak at the2C position (Fig.2b).Among the83EH0plants,all analyzed plants had a signi?cant peak at the2C position similar to that of normal diploids(Fig.2c);however,there were three special EH0plants in which few haploid cells(1C)were detected(Fig.2d),of which two from Zhengdan958and one from F2:3family.Among the 137DH0plants,66plants were detected with large numbers of haploid cells and a few diploid cells (Fig.2e),17plants were detected with equal numbers of haploid and diploid cells(Fig.2f),and the rest had only haploid cells(Table2).The EH0plants were either in a primary diploid state(Fig.2c)or a partial doubled state(Fig.2d).Most of the normal spontane-ously doubled haploid were in a haploid state(Fig.2a) or primary haploid state(Fig.2e),while a few were in a partial doubled state(Fig.2f).

SSR marker analysis

No segments from CAUHOI were detected among the 83EH0plants.The7EH0plants from the Zheng58had

the same genotype as Zheng58,and 17EH 0plants from Chang7-2also had the same genotype as Chang7-2.From all loci characterized in the 33EH 0s from Zhengdan958,47.40%of the alleles were Zheng58-type while 52.60%were Chang7-2-type,producing the expected 1:1ratio for the Zheng58and Chang7-2alleles.The alleles of the rest of EH 0plants from Zhengdan958F 2:3were also from either parent.SSR analysis revealed 100%homozygosis of all the EH 0plants (Table 3;Supplementary Table 2).Agronomic performance of the parents and variations within EH 1lines

The two parents,Zheng58and Chang7-2,greatly varied and differed from each other in the agronomic characteristics being investigated (Table 4).Zheng58differs substantially from Chang7-2in ?ve of the investigated traits such as:plant height,ear height,tassel branch number,ear length and ear width.Highest variation was observed in plant height followed by ear height in the two parents.Chang7-2was taller with the plant height of 188.88cm and the ear height of 83.46cm as compared to Zheng58with 160.00and 59.94cm.The value of EH 1lines and value of mid-parents differed in plant eight,ear height,tassel branch number and ear length,but not statisti-cally signi?cant.The EH 1lines showed a tendency of slightly taller mean height,a little longer ear and less tassel branch number.

No off-type plants were observed within EH 1lines.The homogeneity of within-line populations was further examined by comparing the intra-variance of each EH 1line against the pooled variance of the parents.The F values,determined by the ratio of the intra-variance of the EH 1lines versus the pooled variance of the controls,did not reveal any signi?cant differences in any of the traits tested at a 5%level of signi?cance (Table 4).These indicated that each EH line behaved in the same manner and showed the same potentialities as the parents Chang 7-2and zheng58,which were developed through traditional breeding and evolved through time by ordinary sel?ng.Theo-retically,it is believed that EH is genetically

stable.

Fig.1Morphology of the progenies from Zhengdan9589CAUHOI.a Three types of F 1plants:a1hybrid plant with purple leaves and stem,a2early doubled haploid (EH)plant with green leaves and stalk,a3haploid plant,which was smaller than the diploid.b Selfed ears of fertile plants:b1selfed ears from EH plants,b2selfed ears from normal fertile haploid plants

Table 1Means of normal spontaneously doubled haploids (DH 0)and early spontaneous doubled haploids (EH 0)for various morphological traits

Total

Plant height/cm Ear height/cm Kernel number Ear width/cm Ear length/cm Mean

Mean

Mean Mean Mean DH 045111.45±21.22a 45.89±19.18a 7±1.17a 1.45±0.27a 6.57±1.69a EH 0

33

188.06±24.4b

77.89±15.43b

144±21.33bc

3.78±0.26b

14.09±1.49b

Frequency of early doubled haploids

On the one hand,as for the proportion of EH 0plants among haploid plants (EDR),the average frequencies of con?rmed EH 0plants from Zheng58haploids,Chang7-2haploids,Zhengdan958haploids and the Zhengdan958F 2:3families were 2.24,3.76,3.52and 3.21%,respectively (Table 5).Higher frequency of EH 0plants was detected from Chang7-2than Zheng58.The rate of EH 0occurrence in the Zhengdan958

was

Fig.2Determination of ploidy level using ?ow cytometry according to the position of peaks representing the size of nuclei,as determined for ?ag leaves at the ?owering stage.a Haploids.b Normal diploid plants of the inbred line Zheng58.c Early doubled haploids (EH 0).d EH M ,Mixoploid-type 1was mixoploid in diploid EH plants,with few haploid cells and many diploid cells.e Mixoploid-type 2was chimeric and contained large numbers of haploid cells and few diploid cells.f Mixoploid-type 3had equal amounts of haploid and diploid cells.Note the position of the ?rst peak on the left determines the ploidy.C,haploid cells in G1/G0phase;2C,diploid cells in G1/G0phase and haploid cells in G2/mitosis;4C,diploid cells in G2/mitosis

Table 2Numbers of EH 0and DH 0plants induced from Zhengdan958,the Zhengdan958F 2:3population and the parents with CAUHOI as the inducer line among different ploidy level types,as determined by ?ow cytometry Material/type Total Diploid-type Haploid-type Mixoploid-type 1a Mixoploid-type 2b Mixoploid-type 3c EH 083800300DH 0

137

44

66

17

a Mixoploid-type 1was a chimera with few haploid cells and large numbers of diploid cells

b Mixoploid-type 2was a chimera with large numbers of haploid cells and few diploid cells c

Mixoploid-type 3was a chimera with equal numbers of haploid and diploid cells

between those of the Zheng58and Chang7-2.The average frequency of EH0in these different genetic materials ranged from1to 3.5%(Table5).The frequency of EH in this arti?cially treated population was similar(Table5)to the frequency in spontaneous doubled haploids.On the other hand,however,for the proportion among all of the induced seeds(EHI),the rate was very low.All of the materials’EHI were no more than0.1%(Table5)because of the low HIR of CAUHOI.

Discussion

The production of DH lines makes it easy to carry out genetic studies and signi?cantly shortens the breeding time(Seitz2005).DH technology has become the routine breeding strategy in many commercial maize breeding programs(Geiger and Gordillo2009;Prigge and Melchinger2011).Many works contributed to the haploid induction(Li et al.2009;Wu et al.2014; Zhao et al.2013)but only a few works cared about the haploid doubling.Normally at least two seasons are necessary in the current DH production scheme. One season is haploid induction from manual crossing or in isolation and afterwards haploid kernel identi?-cation depending on special marker such as R1-nj (Chase1969),KOC(Chen and Song2003;Melchinger et al.2013),and so on.The second season is haploid doubling,spontaneously or some chemical agents being used.Theoretically DH1lines can be used for further observation as well as testcross production. However,DH1lines usually cannot be directly used in further tests due to the low number of seeds on selfed ears,hence normally one additional season is required. Recent studies have shown that the mean number of intact seeds on selfed ears(IS)after arti?cial treat-ment was only3.82(Kleiber et al.2012).On the other hand,chemical doubling is time consuming and expensive in many cases.The alternative economic and environmental friendly method of DH produc-tion which can produce enough seed kernels after doubling is very important in DH breeding scheme.

Chalyk(1994)earlier found some‘‘unusual plants phenotypically resembling homozygous lines’’and assumed that the plants resulted from spontaneous chromosome doubling in haploids.By way of expla-nation,he suggested that unusual plants occurred at ‘‘early ontogenetic stages’’.He listed three character-istics of the usual plants:a)they lack heterosis,b)the uniformity of their progeny does not differ from that of inbred lines,and c)they lack dominant marker genes

Table3Frequency of the alleles of parents,heterozygous and inducer in33EH0s generated from the F1of Zhengdan958 Material Heterozygous a Zheng58b Chang7-2c Inducer d

EH01045.6354.370

EH02062.1437.860

EH03052.9447.060

EH04042.0058.000

EH05053.4746.530

EH06050.0050.000

EH07048.9851.020

EH08051.9248.080

EH09062.0038.000

EH010053.8546.150

EH011047.5752.430

EH012060.5839.420

EH013065.0534.950

EH014046.9453.060

EH015048.5451.460

EH016061.3938.610

EH017035.5864.420

EH018064.7135.290

EH019042.3157.690

EH020054.8145.190

EH021053.8546.150

EH022053.4046.600

EH023051.9648.040

EH024054.8145.190

EH025044.1255.880

EH026056.7343.270

EH027055.4544.550

EH028050.6049.400

EH029065.3834.620

EH030053.0646.940

EH031035.5864.420

EH032047.1252.880

EH033063.4636.540

a Frequency of alleles Heterozygous-type in EH

plants through analysis of SSR markers

b Frequency of alleles Zheng58-type,one of the parents,in EH0plants through analysis of SSR markers

c Frequency of alleles Chang7-2-type,one of the parents,in EH0plants through analysis of SSR markers

d Frequency of alleles Inducer-typ

e in EH

plants through analysis of SSR markers

of the pollen parent(Chalyk1994).Chalyk compared the progeny of these plants with?ve-sel?ng inbred lines and found that ear height and ear width of these plants were statistically signi?cant different with inbred lines,so he believed that the inbred lines were quite favorable.And for a long time,no relatively

Table4Mean values for parents(Chang7-2and Zheng58), mid-parent,and early doubled haploid(EH1)lines from Zhengdan958;and F values for variances within EH1lines in comparison with the checks for agronomic traits for six traits (plant height,ear height,tassel length,tassel branch number, ear width and ear length)

Trait Mean F a Signi?cance

of difference Chang7-2Zheng58Mid-parental value EH lines

Plant height/cm188.88±9.60160.00±7.95174.44185.48±8.25b0.05NS

Ear height/cm83.46±8.4659.94±7.0671.777.12±6.900.19NS

Tassel length/cm24.62±3.1824.50±3.2524.5623.63±2.07 1.12NS

Tassel branch number22.31±4.45 6.50±1.4314.4110.16±2.040.61NS

Ear width/cm 4.14±0.41 3.88±0.19 4.01 3.91±0.22 1.31NS

Ear length/cm10.06±0.6314.5±0.1312.2814.16±1.44 2.51NS

NS not signi?cant.Signi?cant at p\0.05

a Ratio of an average of within EH lines variance versus pooled variance of the checks

b An average of standard deviation for each EH

1

line

Table5The frequency of early doubled haploid(EH)plants determined from different materials

Material Total seed Haploid EH HIR%EDR(%)a EHI(%)b

Zhengdan958Beijing35,79971225 1.99 3.510.07 Gansu10,9662278 2.07 3.520.07

A.T.10,6472147 2.01 3.270.07 8701Beijing9,2901573 1.69 1.910.03 Gansu9,2961854 1.99 2.160.04

4F1Beijing7,4101862 2.51 1.080.03 Gansu6,0221622 2.69 1.230.03 Zheng58Beijing6,1091904 3.11 2.110.07 Gansu4,0061273 3.17 2.360.07 Chang7-2Beijing13,1031907 1.45 3.680.05 Gansu16,77426010 1.55 3.850.06

Xu178Beijing6,7161353 2.01 2.220.04 Gansu7,0871462 2.06 1.370.03

Qi319Beijing6,0391253 2.07 2.400.05 Gansu6,3031332 2.11 1.500.03

BY815Beijing9,6852153 2.22 1.400.03 Gansu10,3242233 2.16 1.350.03

B73Beijing11,2323104 2.76 1.290.04 Gansu7,1892023 2.81 1.490.04 Dan598Beijing4,8711892 3.88 1.060.04 Gansu3,5041372 3.91 1.460.06

A.T.arti?cial Treatment(colchicine)

a EDR=(EH plants/total haploid plants in the?eld)9100%

b EHI=(EH plants/total induced seeds)9100%

studies reported the similar phenomenon.In our study, not only cytological observation revealed these plants were diploid plants,but also morphological and molecular analysis of EH revealed that they were as homozygous as normal DH lines and their parents.It revealed homozygosity for100%of the total marker loci in the83EH0plants.The EH1lines from Zhengdan958behaved in the same manner and showed the same potentialities as the parents of Zheng58and Chang7-2(Table4).This was consistent with the results of Chalyk.The comparison between inbred lines and their EH lines showed that the mean values for most of agronomic traits were lower in EH1 lines than in the parental lines(except B73in tassel branch number and8701in ear height and tassel branch number)(Supplementary Table3).This may be from the poorer seed vigor of EH lines or some residual heterozygote loci in inbred lines.However, the differences are not signi?cant.The selfed ears of EH0could produce enough kernels for the next step such as observation and seed production in breeding program(Fig.1b;Table1and Supplementary Fig.1).Our study showed that there was no difference between EH1and DH1lines(Supplementary Table3) in agronomic performance(Sugimoto and Arai2002) for potential use but enough seed kernels of EH can be obtained in one step just after haploid induction. Hence,our research may represents a breakthrough in DH technology of maize research and breeding.

EH0plants were obviously different from DH0 plants in the agronomic performances(Fig.1).This was also re?ected by the degree of diploidization according to?ow cytometry e examination.All of these DH0plants showed incomplete diploidization of the haploid cells whereas the EH0plants showed complete diploidization.The doubling processes between EH and DH are most probably different.On the one hand,in this study,there was little association between the rate of normal spontaneous doubling(data not shown)and the rate of EH.For example,in the line 8,701more than80%of haploids showed fertile pollen and less than3%(EDR)of plants were EH0but in Chang7-2less than30%of haploids were fertile and more than3%(EDR)of plants were EH0 (Table5).On the other hand,the frequency of EH did not increase in conjunction with the increased doubling rate after treatment with a chemical reagent. The fertile haploids produced via chemical(colchi-cine)treatment were also found to be incomplete diploids and the frequency of EH in this arti?cially treated population was similar(Table5)to the frequency in spontaneous doubled haploids.Hence, we speculate that the mechanism of EH formation may be different from the mechanism of normal spontane-ous haploid doubling or that EH formation may happen at a different stage from normal spontaneous haploid doubling but occur by the same mechanism. The arti?cial treatment normally was done during seed germination or in the seedling stage.Even though,the diploidization from arti?cial treatment is not as complete as EH.This implicates that EH formation probably occurs before seed germination or during embryo development after induction pollination.

There are two possible ways that EH plants could arise.One is that maternal materials produce2n female gametes by abnormal meiosis before the induction process and these female gametes develop into diploids by parthenogenesis.The other possibility is that EH can arise from normal n gametes or haploid zygotes produced during the induction by the cross between maternal materials and inducer lines,and in subsequent seed development,the genomes of the embryos were doubled and developed into diploids through abnormal mitosis processes(Mittwoch1978; Segui-Simarro and Nuez2008).Higher plants spon-taneously produce2n gametes at a low frequency(no more than0.6%)(McCoy et al.1982;Lux et al.1990; Bohanec et al.1995;Miyoshi and Asakura1996; Ficcadenti et al.1999);Bauman(1961)found that2n egg cells were produced by abnormal meiosis in a number of maize hybrids at a frequency of0.058%to 0.523%in the crossing of a single cross hybrid with a tetraploid.Many factors can lead to abnormal meiosis that results in the formation of2n gametes,such as meiotic nuclear restitution,second division spindle healing and cytokinesis abnormalities.It seems that this possibility could not be excluded as EHI(the proportion of EH plants among all induced seeds)may be close to frequency of2n gametes frequency reported before.However,all of the planted seeds had obvious R1-nj markers on the top of the embryo and this means that EH could not originate from pathogenesis.And in this study,three mixoploids (EH M)were detected among the EH0plants,in which less than10%of the cells were haploid(Fig.2d).The occurrence of these kinds of mixoploids suggests that the egg or zygote that gave rise to these plants may have been haploid,not diploid,as a few haploid cells

were not completely doubled.Therefore,EHs may have been formed through the second process.

Normally,spontaneous duplication of a haploid genome is thought to occur mainly through three mechanisms:endoreduplication,nuclear fusion and endomitosis(Meyer1925;Hu and Kasha1999; Testillano et al.2004;Shim et al.2006;Segui-Simarro and Nuez2008).If all of these results indicate that EH formation differs from the random formation of2n gametes and represents a special doubling process originating from haploid gametes or zygotes,then the interesting question is whether the haploid comes from a fertilized zygote or an unfertilized maternal gamete. Two mechanisms leading to the formation of maternal haploids have been proposed:(1)one of the two sperm cells that are present during double fertilization is not able to fuse with the egg cell but can trigger haploid embryogenesis.The second sperm cell fuses with the central cell,leading to the formation of a regular triploid endosperm.(2)One of the two sperm cells provided by the inducer is defective but is still able to fuse with the egg cell.During subsequent cell division, the inducer chromosomes degenerate and are elimi-nated stepwise from the primordial endosperm.The second cell fuses with the central cell,as described in the?rst hypothesis(Geiger and Gordillo2009;Li et al. 2009).From our speculation,all spontaneous doubling mechanism might lead to the production of the early doubled haploid.As described above,chromosome elimination could make donor cell chromosomes ‘unstable’,resulting in‘internal division’of the chromosomes themselves combined with a failure in the assembly of the mitotic spindle.This process would happen exactly after normal duplication of the chromatids during S-phase and be induced by the male parent.However,endoreduplication and nuclear fusion need not be exclusive as induction could cause inhibition of mitosis(M-phase)or chromosomes to pass in and out of the cell wall to make it intact because of elimination.

The last but important thing we should mention was that the rate of EH occurrence did not very high:early doubled rate(EDR)was less than4%in our detected materials and this rate was consistent with the results of Chalyk in1994in observing the unusual plants, which was not high enough to reach the reported average spontaneous haploid doubling rate10% (Chase1952;Chalyk1994;Barnabas et al.1999), not to mention the ef?ciency of colchicine treatment;moreover,the other index which Chalyk did not mention was the early doubled haploid induction rate (EHI).Compared to the average haploid induction rate (about2%)of the materials in our study,the EHI was even less.In this situation,EH should be used in plant breeding after further effective studies or technologies to explore to increase the rate.Therefore,taking into account the possible way that the EH plants could arise,we considered that special inducer that could produce enough early doubled haploids would be a new area in further DH breeding.Indeed,the study of EH provides an excellent opportunity to elucidate the mechanisms underlying haploid inducing and dou-bling.Further work is also needed to determine the exact process of spontaneous haploid doubling.Such work will provide new information about the factors that affect double fertilization in?owering plants. Acknowledgments This research was supported by funds from the National863Project(2012AA10A305and2011AA 10A103)and Chinese Universities Scienti?c Fund(15053201). The Zhengdan958F2:3population was generously provided by Dr.Li Haochuan.The authors greatly appreciate the helpful comments and suggestions from Dr.Xu Xiaowei and other anonymous reviewers.

Con?ict of interest The authors declare that they have no con?ict of interest.

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古代晋灵公不君、齐晋鞌之战原文及译文

晋灵公不君(宣公二年) 原文: 晋灵公不君。厚敛以雕墙。从台上弹人,而观其辟丸也。宰夫胹熊蹯不熟,杀之,寘诸畚,使妇人载以过朝。赵盾、士季见其手,问其故而患之。将谏,士季曰:“谏而不入,则莫之继也。会请先,不入,则子继之。”三进及溜,而后视之,曰:“吾知所过矣,将改之。”稽首而对曰:“人谁无过?过而能改,善莫大焉。诗曰:‘靡不有初,鲜克有终。’夫如是,则能补过者鲜矣。君能有终,则社稷之固也,岂惟群臣赖之。又曰:‘衮职有阙,惟仲山甫补之。’能补过也。君能补过,衮不废矣。” 犹不改。宣子骤谏,公患之,使鉏麑贼之。晨往,寝门辟矣,盛服将朝。尚早,坐而假寐。麑退,叹而言曰:“不忘恭敬,民之主也。贼民之主,不忠;弃君之命,不信。有一于此,不如死也!”触槐而死。 秋九月,晋侯饮赵盾酒,伏甲将攻之。其右提弥明知之,趋登曰:“臣侍君宴,过三爵,非礼也。”遂扶以下。公嗾夫獒焉。明搏而杀之。盾曰:“弃人用犬,虽猛何为!”斗且出。提弥明死之。 初,宣子田于首山,舍于翳桑。见灵辄饿,问其病。曰:“不食三日矣!”食之,舍其半。问之,曰:“宦三年矣,未知母之存否。今近焉,请以遗之。”使尽之,而为之箪食与肉,寘诸橐以与之。既而与为公介,倒戟以御公徒,而免之。问何故,对曰:“翳桑之饿人也。”问其名居,不告而退。——遂自亡也。 乙丑,赵穿①攻灵公于桃园。宣子未出山而复。大史书曰:“赵盾弑其君。”以示于朝。宣子曰:“不然。”对曰:“子为正卿,亡不越竟,反不讨贼,非子而谁?”宣子曰:“呜呼!‘我之怀矣,自诒伊戚。’其我之谓矣。” 孔子曰:“董狐,古之良史也,书法不隐。赵宣子,古之良大夫也,为法受恶。惜也,越竞乃免。” 译文: 晋灵公不行君王之道。他向人民收取沉重的税赋以雕饰宫墙。他从高台上用弹弓弹人,然后观赏他们躲避弹丸的样子。他的厨子做熊掌,没有炖熟,晋灵公就把他杀了,把他的尸体装在草筐中,让宫女用车载着经过朝廷。赵盾和士季看到露出来的手臂,询问原由后感到很忧虑。他们准备向晋灵公进谏,士季说:“如果您去进谏而君王不听,那就没有人能够再接着进谏了。还请让我先来吧,不行的话,您再接着来。”士季往前走了三回,行了三回礼,一直到屋檐下,晋灵公才抬头看他。晋灵公说:“我知道我的过错了,我会改过的。”士季叩头回答道:“谁能没有过错呢?有过错而能改掉,这就是最大的善事了。《诗经》说:‘没有人向善没有一个开始的,但却很少有坚持到底的。’如果是这样,那么能弥补过失的人是很少的。您如能坚持向善,那么江山就稳固了,不只是大臣们有所依靠啊。

如何翻译古文

如何翻译古文 学习古代汉语,需要经常把古文译成现代汉语。因为古文今译的过程是加深理解和全面运用古汉语知识解决实际问题的过程,也是综合考察古代汉语水平的过程。学习古代汉语,应该重视古文翻译的训练。 古文翻译的要求一般归纳为信、达、雅三项。“信”是指译文要准确地反映原作的含义,避免曲解原文内容。“达”是指译文应该通顺、晓畅,符合现代汉语语法规范。“信”和“达”是紧密相关的。脱离了“信”而求“达”,不能称为翻译;只求“信”而不顾“达”,也不是好的译文。因此“信”和“达”是文言文翻译的基本要求。“雅”是指译文不仅准确、通顺,而且生动、优美,能再现原作的风格神韵。这是很高的要求,在目前学习阶段,我们只要能做到“信”和“达”就可以了。 做好古文翻译,重要的问题是准确地理解古文,这是翻译的基础。但翻译方法也很重要。这里主要谈谈翻译方法方面的问题。 一、直译和意译 直译和意译是古文今译的两大类型,也是两种不同的今译方法。 1.关于直译。所谓直译,是指紧扣原文,按原文的字词和句子进行对等翻译的今译方法。它要求忠实于原文,一丝不苟,确切表达原意,保持原文的本来面貌。例如: 原文:樊迟请学稼,子曰:“吾不如老农。”请学为圃。子曰:“吾不如老圃。”(《论语?子路》) 译文:樊迟请求学种庄稼。孔子道:“我不如老农民。”又请求学种菜蔬。孔子道:“我不如老菜农。”(杨伯峻《论语译注》) 原文:齐宣王问曰:“汤放桀,武王伐纣,有诸?”(《孟子?梁惠王下》) 译文:齐宣王问道:“商汤流放夏桀,武王讨伐殷纣,真有这回事吗?(杨伯峻《孟子译注》) 上面两段译文紧扣原文,字词落实,句法结构基本上与原文对等,属于直译。 但对直译又不能作简单化理解。由于古今汉语在文字、词汇、语法等方面的差异,今译时对原文作一些适当的调整,是必要的,并不破坏直译。例如: 原文:逐之,三周华不注。(《齐晋鞌之战》) 译文:〔晋军〕追赶齐军,围着华不注山绕了三圈。

齐晋鞌之战原文和译文

鞌之战选自《左传》又名《鞍之战》原文:楚癸酉,师陈于鞌(1)。邴夏御侯,逢丑父为右②。晋解张御克,郑丘缓为右(3)。侯日:“余姑翦灭此而朝食(4)”。不介马而驰之⑤。克伤于矢,流血及屦2 未尽∧6),曰:“余病矣(7)!”张侯曰:“自始合(8),而矢贯余手及肘(9),余折以御,左轮朱殷(10),岂敢言病吾子忍之!”缓曰:“自始合,苟有险,余必下推车,子岂_识之(11)然子病矣!”张侯曰:“师之耳目,在吾旗鼓,进退从之。此车一人殿之(12),可以集事(13),若之何其以病败君之大事也擐甲执兵(14),固即死也(15);病未及死,吾子勉之(16)!”左并辔(17) ,右援拐鼓(18)。马逸不能止(19),师从之,师败绩。逐之,三周华不注(20) 韩厥梦子舆谓己曰:“旦辟左右!”故中御而从齐侯。邴夏曰:“射其御者,君子也。”公曰:“谓之君子而射之,非礼也。”射其左,越于车下;射其右,毙于车中。綦毋张丧车,从韩厥,曰:“请寓乘。”从左右,皆肘之,使立于后。韩厥俛,定其右。逢丑父与公易位。将及华泉,骖絓于木而止。丑父寝于轏中,蛇出于其下,以肱击之,伤而匿之,故不能推车而及。韩厥执絷马前,再拜稽首,奉觞加璧以进,曰:“寡君使群臣为鲁、卫请,曰:‘无令舆师陷入君地。’下臣不幸,属当戎行,无所逃隐。且惧奔辟而忝两君,臣辱戎士,敢告不敏,摄官承乏。” 丑父使公下,如华泉取饮。郑周父御佐车,宛茷为右,载齐侯以免。韩厥献丑父,郤献子将戮之。呼曰:“自今无有代其君任患者,有一于此,将为戮乎”郤子曰:“人不难以死免其君,我戮之不祥。赦之,以劝事君者。”乃免之。译文1:在癸酉这天,双方的军队在鞌这个地方摆开了阵势。齐国一方是邴夏为齐侯赶车,逢丑父当车右。晋军一方是解张为主帅郤克赶车,郑丘缓当车右。齐侯说:“我姑且消灭了这些人再吃早饭。”不给马披甲就冲向了晋军。郤克被箭射伤,血流到了鞋上,但是仍不停止擂鼓继续指挥战斗。他说:“我受重伤了。”解张说:“从一开始接战,一只箭就射穿了我的手和肘,左边的车轮都被我的血染成了黑红色,我哪敢说受伤您忍着点吧!”郑丘缓说:“从一开始接战,如果遇到道路不平的地方,我必定(冒着生命危险)下去推车,您难道了解这些吗不过,您真是受重伤了。”daier 解张说:“军队的耳朵和眼睛,都集中在我们的战旗和鼓声,前进后退都要听从它。这辆车上还有一个人镇守住它,战事就可以成功。为什么为了伤痛而败坏国君的大事呢身披盔甲,手执武器,本来就是去走向死亡,伤痛还没到死的地步,您还是尽力而为吧。”一边说,一边用左手把右手的缰绳攥在一起,用空出的右手抓过郤克手中的鼓棰就擂起鼓来。(由于一手控马,)马飞快奔跑而不能停止,晋军队伍跟着指挥车冲上去,把齐军打得打败。晋军随即追赶齐军,三次围绕着华不注山奔跑。韩厥梦见他去世的父亲对他说:“明天早晨作战时要避开战车左边和右边的位置。”因此韩厥就站在中间担任赶车的来追赶齐侯的战车。邴夏说:“射那个赶车的,他是个君子。”齐侯说: “称他为君子却又去射他,这不合于礼。”daier 于是射车左,车左中箭掉下了车。又射右边的,车右也中箭倒在了车里。(晋军的)将军綦毋张损坏了自己的战车,跟在韩厥的车后说: “请允许我搭乗你的战车。”他上车后,无论是站在车的左边,还是站在车的右边,韩厥都用肘推他,让他站在自己身后——战车的中间。韩厥又低下头安定了一下受伤倒在车中的那位自己的车右。于是逢丑父和齐侯(乘韩厥低头之机)互相调换了位置。将要到达华泉时,齐侯战车的骖马被树木绊住而不能继续逃跑而停了下来。(头天晚上)逢丑父睡在栈车里,有一条蛇从他身子底下爬出来,他用小臂去打蛇,小臂受伤,但他(为了能当车右)隐瞒了这件事。由于这样,他不能用臂推车前进,因而被韩厥追上了。韩厥拿着拴马绳走到齐侯的马前,两次下拜并行稽首礼,捧着一杯酒并加上一块玉璧给齐侯送上去,

《鞌之战》阅读答案(附翻译)原文及翻译

《鞌之战》阅读答案(附翻译)原文及翻 译 鞌之战[1] 选自《左传成公二年(即公元前589年)》 【原文】 癸酉,师陈于鞌[2]。邴夏御齐侯[3],逢丑父为右[4]。晋解张御郤克,郑丘缓为右[5]。齐侯曰:余姑翦灭此而朝食[6]。不介马而驰之[7]。郤克伤于矢,流血及屦,未绝鼓音[8],曰:余病[9]矣!张侯[10]曰:自始合,而矢贯余手及肘[11],余折以御,左轮朱殷[12],岂敢言病。吾子[13]忍之!缓曰:自始合,苟有险[14],余必下推车,子岂识之[15]?然子病矣!张侯曰:师之耳目,在吾旗鼓,进退从之[16]。此车一人殿之[17],可以集事[18],若之何其以病败君之大事也[19]?擐甲执兵,固即死也[20]。病未及死,吾子勉之[21]!左并辔[22],右援枹而鼓[23],马逸不能止[24],师从之。齐师败绩[25]。逐之,三周华不注[26]。 【注释】 [1]鞌之战:春秋时期的著名战役之一。战争的实质是齐、晋争霸。由于齐侯骄傲轻敌,而晋军同仇敌忾、士气旺盛,战役以齐败晋胜而告终。鞌:通鞍,齐国地名,在今山东济南西北。 [2]癸酉:成公二年的六月十七日。师,指齐晋两国军队。陈,

列阵,摆开阵势。 [3]邴夏:齐国大夫。御,动词,驾车。御齐侯,给齐侯驾车。齐侯,齐国国君,指齐顷公。 [4]逢丑父:齐国大夫。右:车右。 [5]解张、郑丘缓:都是晋臣,郑丘是复姓。郤(x )克,晋国大夫,是这次战争中晋军的主帅。又称郤献子、郤子等。 [6]姑:副词,姑且。翦灭:消灭,灭掉。朝食:早饭。这里是吃早饭的意思。这句话是成语灭此朝食的出处。 [7]不介马:不给马披甲。介:甲。这里用作动词,披甲。驰之:驱马追击敌人。之:代词,指晋军。 [8] 未绝鼓音:鼓声不断。古代车战,主帅居中,亲掌旗鼓,指挥军队。兵以鼓进,击鼓是进军的号令。 [9] 病:负伤。 [10]张侯,即解张。张是字,侯是名,人名、字连用,先字后名。 [11]合:交战。贯:穿。肘:胳膊。 [12]朱:大红色。殷:深红色、黑红色。 [13]吾子:您,尊敬。比说子更亲切。 [14]苟:连词,表示假设。险:险阻,指难走的路。 [15]识:知道。之,代词,代苟有险,余必下推车这件事,可不译。 [16]师之耳目:军队的耳、目(指注意力)。在吾旗鼓:在我们

《鞌之战》阅读答案附翻译

《鞌之战》阅读答案(附翻译) 《鞌之战》阅读答案(附翻译) 鞌之战[1] 选自《左传·成公二年(即公元前589年)》 【原文】 癸酉,师陈于鞌[2]。邴夏御齐侯[3],逢丑父为右[4]。晋解张御郤克,郑丘缓为右[5]。齐侯曰:“余姑 翦灭此而朝食[6]。”不介马而驰之[7]。郤克伤于矢, 流血及屦,未绝鼓音[8],曰:“余病[9]矣!”张侯[10]曰:“自始合,而矢贯余手及肘[11],余折以御,左轮 朱殷[12],岂敢言病。吾子[13]忍之!”缓曰:“自始合,苟有险[14],余必下推车,子岂识之[15]?——然 子病矣!”张侯曰:“师之耳目,在吾旗鼓,进退从之[16]。此车一人殿之[17],可以集事[18],若之何其以 病败君之大事也[19]?擐甲执兵,固即死也[20]。病未 及死,吾子勉之[21]!”左并辔[22],右援枹而鼓[23],马逸不能止[24],师从之。齐师败绩[25]。逐之,三周 华不注[26]。 【注释】 [1]鞌之战:春秋时期的著名战役之一。战争的实质是齐、晋争霸。由于齐侯骄傲轻敌,而晋军同仇敌忾、 士气旺盛,战役以齐败晋胜而告终。鞌:通“鞍”,齐

国地名,在今山东济南西北。 [2]癸酉:成公二年的六月十七日。师,指齐晋两国军队。陈,列阵,摆开阵势。 [3]邴夏:齐国大夫。御,动词,驾车。御齐侯,给齐侯驾车。齐侯,齐国国君,指齐顷公。 [4]逢丑父:齐国大夫。右:车右。 [5]解张、郑丘缓:都是晋臣,“郑丘”是复姓。郤(xì)克,晋国大夫,是这次战争中晋军的主帅。又称郤献子、郤子等。 [6]姑:副词,姑且。翦灭:消灭,灭掉。朝食:早饭。这里是“吃早饭”的意思。这句话是成语“灭此朝食”的出处。 [7]不介马:不给马披甲。介:甲。这里用作动词,披甲。驰之:驱马追击敌人。之:代词,指晋军。 [8]未绝鼓音:鼓声不断。古代车战,主帅居中,亲掌旗鼓,指挥军队。“兵以鼓进”,击鼓是进军的号令。 [9]病:负伤。 [10]张侯,即解张。“张”是字,“侯”是名,人名、字连用,先字后名。 [11]合:交战。贯:穿。肘:胳膊。 [12]朱:大红色。殷:深红色、黑红色。 [13]吾子:您,尊敬。比说“子”更亲切。

左传《齐晋鞌之战》原文+翻译+注释

左传《齐晋鞌之战》原文+翻译+注释 楚癸酉,师陈于鞌(1)。邴夏御侯,逢丑父为右②。晋解张御克,郑丘缓 为右(3)。侯日:“余姑翦灭此而朝食(4)”。不介马而驰之⑤。克伤于矢, 流血及屦2未尽∧?6),曰:“余病矣(7)!”张侯曰:“自始合(8),而矢贯余手 及肘(9),余折以御,左轮朱殷(10),岂敢言病?吾子忍之!”缓曰:“自始合,苟有险,余必下推车,子岂_识之(11)?然子病矣!”张侯曰:“师之耳目,在 吾旗鼓,进退从之。此车一人殿之(12),可以集事(13),若之何其以病败君之大事也?擐甲执兵(14),固即死也(15);病未及死,吾子勉之(16)!”左并辔(17) ,右援拐?鼓(18)。马逸不能止(19),师从之,师败绩。逐之,三周华不注(20) 韩厥梦子舆谓己曰:“旦辟左右!”故中御而从齐侯。邴夏曰:“射其御者,君子也。”公曰:“谓之君子而射之,非礼也。”射其左,越于车下;射其右,毙于车中。綦毋张丧车,从韩厥,曰:“请寓乘。”从左右,皆肘之,使立于后。韩厥俛,定其右。逢丑父与公易位。将及华泉,骖絓于木而止。丑父寝于轏中,蛇出于其下,以肱击之,伤而匿之,故不能推车而及。韩厥执絷马前,再拜稽首,奉觞加璧以进,曰:“寡君使群臣为鲁、卫请,曰:‘无令舆师陷入君地。’下臣不幸,属当戎行,无所逃隐。且惧奔辟而忝两君,臣辱戎士,敢告不敏,摄官承乏。”丑父使公下,如华泉取饮。郑周父御佐车,宛茷为右,载齐侯以免。韩厥献丑父,郤献子将戮之。呼曰:“自今无有代其君任患者,有一于此,将为戮乎?”郤子曰:“人不难以死免其君,我戮之不祥。赦之,以劝事君者。”乃免之。 在癸酉这天,双方的军队在鞌这个地方摆开了阵势。齐国一方是邴夏为齐侯赶车,逢丑父当车右。晋军一方是解张为主帅郤克赶车,郑丘缓当车右。齐侯说:“我姑且消灭了这些人再吃早饭。”不给马披甲就冲向了晋军。郤克被箭射伤,血流到了鞋上,但是仍不停止擂鼓继续指挥战斗。他说:“我受重伤了。”解张说:“从一开始接战,一只箭就射穿了我的手和肘,左边的车轮都被我的血染成了黑红色,我哪敢说受伤?您忍着点吧!”郑丘缓说:“从一开始接战,如果遇到道路不平的地方,我必定(冒着生命危险)下去推车,您难道了解这些吗?不过,您真是受重伤了。”daier解张说:“军队的耳朵和眼睛,都集中在我们的战旗和鼓声,前进后退都要听从它。这辆车上还有一个人镇守住它,战事就可以成功。为什么为了伤痛而败坏国君的大事呢?身披盔甲,手执武器,本来就是去走向死亡,伤痛还没到死的地步,您还是尽力而为吧。”一边说,一边用左手把右手的缰绳攥在一起,用空出的右手抓过郤克手中的鼓棰就擂起鼓来。(由于一手控马,)马飞快奔跑而不能停止,晋军队伍跟着指挥车冲上去,把齐军打得打败。晋军随即追赶齐军,三次围绕着华不注山奔跑。

《鞌之战》阅读答案(附翻译)

鞌之战[1]选自《左传·成公二年(即公元前589年)》【原文】癸酉,师陈于鞌[2]。邴夏御齐侯[3],逢丑父为右[4]。晋解张御郤克,郑丘缓为右[5]。齐侯曰:“余姑翦灭此而朝食[6]。”不介马而驰之[7]。郤克伤于矢,流血及屦,未绝鼓音[8],曰:“余病[9]矣!”张侯[10]曰:“自始合,而矢贯余手及肘[11],余折以御,左轮朱殷[12],岂敢言病。吾子[13]忍之!”缓曰:“自始合,苟有险[14],余必下推车,子岂识之[15]?——然子病矣!”张侯曰:“师之耳目,在吾旗鼓,进退从之[16]。此车一人殿之[17],可以集事[18],若之何其以病败君之大事也[19]?擐甲执兵,固即死也[20]。病未及死,吾子勉之[21]!”左并辔[22],右援枹而鼓[23],马逸不能止[24],师从之。齐师败绩[25]。逐之,三周华不注[26]。【注释】 [1]鞌之战:春秋时期的著名战役之一。战争的实质是齐、晋争霸。由于齐侯骄傲轻敌,而晋军同仇敌忾、士气旺盛,战役以齐败晋胜而告终。鞌:通“鞍”,齐国地名,在今山东济南西北。 [2]癸酉:成公二年的六月十七日。师,指齐晋两国军队。陈,列阵,摆开阵势。 [3]邴夏:齐国大夫。御,动词,驾车。御齐侯,给齐侯驾车。齐侯,齐国国君,指齐顷公。 [4]逢丑父:齐国大夫。右:车右。 [5]解张、郑丘缓:都是晋臣,“郑丘”是复姓。郤(xì)克,晋国大夫,是这次战争中晋军的主帅。又称郤献子、郤子等。 [6]姑:副词,姑且。翦灭:消灭,灭掉。朝食:早饭。这里是“吃早饭”的意思。这句话是成语“灭此朝食”的出处。 [7]不介马:不给马披甲。介:甲。这里用作动词,披甲。驰之:驱马追击敌人。之:代词,指晋军。 [8] 未绝鼓音:鼓声不断。古代车战,主帅居中,亲掌旗鼓,指挥军队。“兵以鼓进”,击鼓是进军的号令。 [9] 病:负伤。 [10]张侯,即解张。“张”是字,“侯”是名,人名、字连用,先字后名。 [11]合:交战。贯:穿。肘:胳膊。 [12]朱:大红色。殷:深红色、黑红色。 [13]吾子:您,尊敬。比说“子”更亲切。 [14]苟:连词,表示假设。险:险阻,指难走的路。 [15]识:知道。之,代词,代“苟有险,余必下推车”这件事,可不译。 [16]师之耳目:军队的耳、目(指注意力)。在吾旗鼓:在我们的旗子和鼓声上。进退从之:前进、后退都听从它们。 [17]殿之:镇守它。殿:镇守。 [18]可以集事:可以(之)集事,可以靠它(主帅的车)成事。集事:成事,指战事成功。 [19]若之何:固定格式,一般相当于“对……怎么办”“怎么办”。这里是和语助词“其”配合,放在谓语动词前加强反问,相当于“怎么”“怎么能”。以,介词,因为。败,坏,毁坏。君,国君。大事,感情。古代国家大事有两件:祭祀与战争。这里指战争。 [20]擐:穿上。执兵,拿起武器。 [21]勉,努力。 [22]并,动词,合并。辔(pèi):马缰绳。古代一般是四匹马拉一车,共八条马缰绳,两边的两条系在车上,六条在御者手中,御者双手执之。“左并辔”是说解张把马缰绳全合并到左手里握着。 [23]援:拿过来。枹(fú):击鼓槌。鼓:动词,敲鼓。 [24]逸:奔跑,狂奔。 [25] 败绩:大败。 [26] 周:环绕。华不注:山名,在今山东济南东北。【译文】六月十七日,齐晋两军在鞌地摆开阵势。邴夏为齐侯驾车,逢丑父担任车右做齐侯的护卫。晋军解张替郤克驾车,郑丘缓做了郤克的护卫。齐侯说:“我姑且消灭了晋军再吃早饭!”齐军没有给马披甲就驱车进击晋军。郤克被箭射伤,血一直流到鞋上,但他一直没有停止击鼓进。并说:“我受重伤了!”解张说:“从开始交战,箭就射穿了我的手和胳膊肘,我折断箭杆继续驾车,左边的车轮被血染得深红色,哪里敢说受了重伤?您还是忍住吧。”郑丘缓说:“从开始交战,只要遇到险峻难走的路,我必定要下去推车,您哪里知道这种情况呢?——不过您确实受重伤了!”解张说:“我们的旗帜和战鼓是军队的耳目,或进或退都听从旗鼓指挥。这辆战车只要一人镇守,就可以凭它成事。怎么能因为受伤而败坏国君的大事呢?穿上铠甲,拿起武器,本来就抱定了必死的决心。您虽然受了重伤还没有到死的地步,您就尽最大的努力啊!”于是左手把马缰绳全部握在一起,右手取过鼓槌来击鼓。战马狂奔不止,晋军跟着主帅的车前进。齐军大败,晋军追击齐军,绕着华不注山追了三圈。

齐晋鞌之战的全文翻译

齐晋鞌之战的全文翻译 1、版本一 晋鞌之战(成公二年) 作者:左丘明 【原文】 楚癸酉,师陈于鞌(1)。邴夏御侯,逢丑父为右②。晋解张御克,郑丘缓为右(3)。侯日:“余姑翦灭此而朝食(4)”。不介马而驰之⑤。克伤于矢,流血及屨2未尽∧?6),曰:“余病矣(7)!”张侯曰:“自始合(8),而矢贯余手及肘(9),余折以御,左轮朱殷(10),岂敢言病?吾子忍之!”缓曰:“自始合,苟有险,余必下推车,子岂_识之(11)?然子病矣!”张侯曰:“师之耳目,在吾旗鼓,进退从之。此车一人殿之(12),可以集事(13),若之何其以病败君之大事也?擐甲执兵(14),固即死也(15);病未及死,吾子勉之(16)!”左并辔(17),右援枴?鼓(18)。马逸不能止(19),师从之,师败绩。逐之,三周华不注(20)。 【注释】 ①师:指、晋两国军队。羞:同“鞍”,国地名,在今山东济南西北。②邴(bing)夏:国大夫。侯:顷公。逢丑父:国大夫。右:车右。③解张:晋国大夫,又称张侯。克:即献子,晋国大大,晋军主帅。郑丘缓:晋国大夫,姓郑丘,名缓。(4)姑:暂且。翦灭:消灭。此;指晋军。朝食;吃早饭。⑤不介马:不给马披甲。驰之:驱马追击敌人。(6)未绝鼓音:作战时,主帅亲自掌旗鼓,指挥三军,

所以克受伤后仍然击鼓不停。(7)病:负伤。(8)合:交战。(9)贯:射。穿。肘:胳膊。(10)朱:大红色。殷:深红色。(11)识:知道。 (12) 殿:镇守。(13)集事:成事。(14)擐(huan):穿上。兵:武器。 (15) 即:就。即死:就死,赴死。(16)勉:努力。(17)并:合在一起。辔(Pei):马组绳。(18)援:拉过来。枴〉襲):鼓槌。(19)逸:奔跑,狂奔。(20)周:环绕华不注:山名,在今山东济南东北。 【译文】 六月十七日,国和晋国的军队在鞌摆开了阵势。邴夏为顷公驾车,逢丑父担任车右。晋国解张为卻克驾车,郑丘缓担任车右。顷公说:“我暂且先消灭了这些敌人再吃早饭。”军没有给马披甲就驱车进击晋军。卻克被箭射伤,血流到鞋子上,但他一直没有停止击鼓,并说:“我受伤了!”解张说:“从开始交战,我的手和胳膊就被箭射穿了,我折断了箭,继续驾车,左边的车轮因被血染成了深红色,哪里敢说受了伤?您还是忍住吧?”郑丘缓说:“从开始交战,只要遇到险阻,我一定要下去推车,您哪里知道这些?可是您却受伤了!”解张说:“我们的旗帜和战鼓是军队的耳目,军队进攻和后撤都听从旗鼓指挥。这辆战车只要一个人镇守,就可以成功,怎么能因为负了伤而败坏国君的大事呢?穿上铠甲,拿起武器,本来就是去赴死;受伤不到死的地步,您要奋力而为啊!”解张左手把马绳全部握在一起,右手拿过鼓槌来击鼓。战马狂奔不已,晋军跟著主帅的车前进,军大败,晋军追击军,围著华不注山追了三圈。 2、版本二

齐晋鞌之战原文和译文

创作编号:BG7531400019813488897SX 创作者:别如克* 鞌之战选自《左传》又名《鞍之战》原文:楚癸酉,师陈于鞌(1)。邴夏御侯,逢丑父为右②。晋解张御克,郑丘缓为右(3)。侯日:“余姑翦灭此而朝食(4)”。不介马而驰之⑤。克伤于矢,流血及屦2 未尽∧?6),曰:“余病矣(7)!”张侯曰:“自始合(8),而矢贯余手及肘(9),余折以御,左轮朱殷(10),岂敢言病?吾子忍之!”缓曰:“自始合,苟有险,余必下推车,子岂_识之(11)?然子病矣!”张侯曰:“师之耳目,在吾旗鼓,进退从之。此车一人殿之(12),可以集事(13),若之何其以病败君之大事也?擐甲执兵(14),固即死也(15);病未及死,吾子勉之(16)!”左并辔(17) ,右援拐?鼓(18)。马逸不能止(19),师从之,师败绩。逐之,三周华不注(20) 韩厥梦子舆谓己曰:“旦辟左右!”故中御而从齐侯。邴夏曰:“射其御者,君子也。”公曰:“谓之君子而射之,非礼也。”射其左,越于车下;射其右,毙于车中。綦毋张丧车,从韩厥,曰:“请寓乘。”从左右,皆肘之,使立于后。韩厥俛,定其右。逢丑父与公易位。将及华泉,骖絓于木而止。丑父寝于轏中,蛇出于其下,以肱击之,伤而匿之,故不能推车而及。韩厥执絷马前,再拜稽首,奉觞加璧以进,曰:“寡君使群臣为鲁、卫请,曰:‘无令舆师陷入君地。’下臣不幸,属当戎行,无所逃隐。且惧奔辟而忝两君,臣辱戎士,敢告不敏,摄官承乏。” 丑父使公下,如华泉取饮。郑周父御佐车,宛茷为右,载齐侯以免。韩厥献丑父,郤献子将戮之。呼曰:“自今无有代其君任患者,有一于此,将为戮乎?”郤子曰:“人不难以死免其君,我戮之不祥。赦之,以劝事君者。”乃免之。译文1:在癸酉这天,双方的军队在鞌这个地方摆开了阵势。齐国一方是邴夏为齐侯赶车,逢丑父当车右。晋军一方是解张为主帅郤克赶车,郑丘缓当车右。齐侯说:“我姑且消灭了这些人再吃早饭。”不给马披甲就冲向了晋军。郤克被箭射伤,血流到了鞋上,但是仍不停止擂鼓继续指挥战斗。他说:“我受重伤了。”解张说:“从一开始接战,一只箭就射穿了我的手和肘,左边的车轮都被我的血染成了黑红色,我哪敢说受伤?您忍着点吧!”郑丘缓说:“从一开始接战,如果遇到道路不平的地方,我必定(冒着生命危险)下去推车,您难道了解这些吗?不过,您真是受重伤了。”daier 解张说:“军队的耳朵和眼睛,都集中在我们的战旗和鼓声,前进后退都要听从它。这辆车上还有一个人镇守住它,战事就可以成功。为什么为了伤痛而败坏国君的大事呢?身披盔甲,手执武器,本来就是去走向死亡,伤痛还没到死的地步,您还是尽力而为吧。”一边说,一边用左手把右手的缰绳攥在一起,用空出的右手抓过郤克手中的鼓棰就擂起鼓来。(由于一手控马,)马飞快奔跑而不能停止,晋军队伍跟着指挥车冲上去,把齐军打得打败。晋军随即追赶齐军,三次围绕着华不注山奔跑。韩厥梦见他去世的父亲对他说:“明天早晨作战时要避开

齐晋鞌之战(原文和译文)

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