The crossing of common wheat (Triticum aestivum L.) with cultivated rye (Secale cereale L.). I. crossability,pollen grain germination and pollen tube growth

Summary The crossing of common wheat (Triticum aestivum L.) with rye (Secale cereale L.), and especially the action of the crossability genes of wheat, was studied using the readily crossable wheat cv Chinese Spring (genotype kr ₁ kr ₁ kr ₂ kr ₂ >), the poorly crossable wheat cv Hope (genotype Kr ₁ Kr ₁ Kr ₂ Kr ₂ ), as well as the disomic substitution line of chromosome 5B of Hope into Chinese Spring (CS/Hope 5B, genotype Kr ₁ Kr ₁ kr ₂ kr ₂ ). By comparing crossability and actual fertilization, the poor crossability with rye of both cv Hope and the CS/Hope 5B substitution line was shown to result from absence of fertilization. Studies of pollen grain germination and pollen tube growth showed that the dominant alleles of the crossability genes manifested themselves through retardation and eventually inhibition of pollen tube growth at the style base and in the ovary wall. In Hope the growth of all pollen tubes was inhibited, whereas in CS/Hope 5B rarely fertilization was achieved. The recessive alleles of the crossability genes do not seem to have an influence on the growth of rye pollen tubes in wheat pistils.     

The crossing of common wheat (Triticum aestivum L.) with cultivated rye (Secale cereale L.). II. Fertilization and early post-fertilization developments

Summary Fertilization and early seed development was studied in the variety Chinese Spring of common wheat (Triticum aestivum L.) after pollination with rye (Secale cereale L.) and selfing, and in the common wheat variety Hope after selfing. In all three combinations the first pollen tube reached the micropyle in about 40 min after pollination. When pollinated with rye the migration of the sperm nuclei to the egg cell and the polar nuclei was delayed by about an hour. In the subsequent development until 72 hours after pollination the average cellular and nuclear doubling times of embryo and endosperm were about 16 and 8 hours for the interspecific combination. 18 and 9 hours for Chinese Spring selfed and 20 and 12 hours for Hope selfed.     

The site of action of crossability genes (Kr 1, Kr 2) between Triticum and Secale. II. Proportion of pistils containing pollen tubes and effects of alternate pollination on seed set

Summary Different wheat genotypes (T. aestivum) were crossed with rye to ascertain the site or sites of manifestation of the crossability genes, Kr ₁ and Kr ₂. By using fluorescence microscopy, it was found that the order of increasing proportion of wheat micropyles containing pollen tubes is strongly correlated with the levels of crossability with rye. High crossable genotypes have more micropyles containing pollen tubes than those of the low crossable ones. Most of the inhibition or retardation of pollen tubes occurred between the style base and top of the embryo sac, expecially with the low crossable genotypes where both Kr-genes are present. The results also indicate that Kr ₁ is a greater inhibitor than Kr ₂. Seed set is also highly correlated with the number or proportion of micropyles having pollen tubes. Alternate pollinations seem to support the view that rye pollen tubes do not reach the micropyles of the low crossable genotypes, and hence when repollinated with wheat selfed seeds are produced.     

Factors affecting haploid production in wheat using the Hordeum bulbosum system. 1. Genotypic and environmental effects on pollen grain germination,pollen tube growth and the frequency of fertilization

Summary A quantitative examination of pollen grain germination and pollen tube growth within the overy wall was made in selected wheat × tetraploid H. bulbosum hybridizations, to investigate the effect of the wheat and the H. bulbosum genotype on these characters. As expected, variation at the known crossability loci had no effect on pollen grain germination. The frequency of pollen tube penetration of the ovary wall was, however, severely reduced when the dominant alleles were present. Pollen tube penetration was nevertheless observed in 3 of the 110 ovaries examined of the non-crossable cultivar Highbury. The H. bulbosum genotype had a much smaller effect on these characters, but significant differences between the clones were observed in the numbers of pollen tubes initially penetrating the ovary wall. Although two H. bulbosum genotypes showed no significant differences in the number of ovaries with pollen tubes at the base of the ovule, significant differences in the frequency of fertilization were observed. The possible cause of this discrepancy is discussed.The frequency of fertilization in crossable wheat × H. bulbosum hybridizations was improved by the application of gibberellic acid within 10 minutes of pollination, and reduced by an increase in the ambient temperature from 20°C to 26°C. Fertilization following the pollination of non-crossable wheat genotypes was not affected by either of these factors.     

Transfer of the recessive crossability allele kr1 from Chinese Spring into the winter wheat variety Martonvásári 9

Summary The recessive of crossability allele kr1 was transferred from the spring wheat variety Chinese Spring (CS) into the winter wheat variety Martonvásári 9 (Mv9) by backcrossing the Mv9 × CS hybrids with Mv9. The Mv9 variety possesses dominant Kr1 alleles and is heterogeneous at the kr2 locus, so that some individual plants carry recessive kr2 alleles. The selection of plants possessing the recessive kr alleles from the (Mv9 × CS)Mv9 BC₁ generation was carried out according to the seed set achieved when pollinated with rye (Secale cereale L. cv. Mercator). The partial dominance of the Kr alleles made it possible to differentiate between plants heterozygous at the Kr1 locus and Kr1Kr1 homozygous dominant plants. Two selfed consecutive progenies were tested by pollination with rye to select the homozygous recessive kr1kr1kr2kr2 plants and to check the result of the selection after each backcross.As a result of three backcrosses with Mv9 and two selfings after each backcross the selected progenies had 61.6% seed set with rye tested on sixty individual plants. These data confirm that after the third backcross the selected Mv9 kr1 line carries necessive crossability alleles Kr1 and Kr2, but the genotype is 93.75% Mv9.     

The site of action of the crossability genes (Kr 1, Kr 2) between Triticum and Secale. I. Pollen germination,pollen tube growth,and number of pollen tubes

Summary Seven genotypes of common wheat (Triticum aestivum L.) were crossed with rye (Secale cereale L.) in order to find the site or sites of action of the crossability genes, Kr ₁ and Kr ₂, of wheat. The data obtained, by fluorescence microscopy, were compared to the controls (wheat x wheat). The results indicate that the crossability genes have little effect on pollen germination and on the time taken for the pollen tubes to reach the microphyle, irrespective of their crossabilities with rye. The number of pollen tubes reaching the microphyle is, however, affected by the Kr-genes, as high crossable genotypes have more pollen tubes than the low crossable ones. There was a high correlation between the mean number of pollen tubes at the micropyle with seed set, which also reflects the crossability. The Kr-genes seem to manifest themselves in the retardation and inhibition of pollen tube growth between the style base and the top of the embryo sac, where the effect is most distinct in the low crossable genotypes.     

Factors affecting haploid production in wheat using the Hordeum bulbosum system. 2. The effect of the timing of pollination

Summary The effect of manipulating the timing of pollination on the fertilization frequency in wheat × tetraploid Hordeum bulbosum hybridizations was investigated as a possible means of overcoming varietal incompatibility in this cross. Pre- and post-anthesis pollinations were made on previously identified highly crossable and poorly crossable wheat genotypes and the stage of development of the individual florets at pollination was expressed in days from anthesis units.In both crossable and non-crossable wheat genotypes, a significant negative linear relationship was observed between floret age and fertilization frequency. The magnitude of the response of early pollination was dependent on the number of alleles for non-crossability present at the Kr loci. The non-crossable cultivar Highbury, possessing more than one allele for incompatibility, showed a slight response. The greatest response was shown by the highly crossable cultivar Chinese Spring, possessing alleles for crossability at all Kr loci. The single chromosome substitution line, Chinese Spring (Hope 5B), possessing the most potent allele for non-crossability, Kr ₁, showed an intermediate response. The H. bulbosum genotype had no effect on the magnitude of this response, influencing only the mean fertilization frequency.     

Introgression of Dasypyrum villosum chromatin into common wheat improves grain protein quality

Dasypyrum villosum (L.) Candargy (DV) is adiploid (2n = 14, VV genomes), allogamous grass of theMediterranean region. It may be hybridized with wheatand is thus a gene resource for wheat improvement. Westudied grain protein concentration andSDS-sedimentation (SED) as indicators of end-usequality. The latter is a good predictor of glutenstrength. A-PAGE and SDS-PAGE were used to identifymonomeric and polymeric seed storage proteins,respectively, to relate proteins of DV to those foundin Chinese Spring (CS), Triticum aestivum L.,wheat. Two full-sib lines of DV had high grain protein(19.3 and 20.3%), but one had very low mean SED (69mm) and one had very high (118 mm) based on onegreenhouse and one field test. CS had very low grainprotein (12.0%) and weak gluten (33 mm). Single-DVchromosome addition and substitution lines and twoDV-wheat recombinant lines all had higher grainprotein than CS (range 13.9 to 16.7%). SED valuesshowed a different pattern. CS=4V and CS=6V hadlow SED, 63 and 44 mm, similar to CS, whereas CS=1Vand full sib DV 200 had very strong gluten, 118 mm, asdid substitution lines CS1V (1A) and CS1V (1B), 125and 131 mm, respectively. One hybrid-derived line withDV-wheat 1V recombinant chromosome had SED of 99 mmand one line with a 6V added chromosome had SED of 64mm. The large positive effects of quality in the wheathaving DV chromosome 1V are believed to be due to DValleles at the Glu-V1 and Gli-V1/Glu-V3loci. DV chromosomes 4V and 6V did not contribute toimproved quality probably due to Gli-V2 and Gli-V3 which, as the orthologous loci in wheat, donot enhance wheat quality. Based on the positiveeffects of alleles on DV chromosome 1V in a breadwheat background, we conclude that D. villosumis a source of allelic diversity that can beconsidered for improving end-use quality in breadwheat.     

Temperature as a component of the expression of developmental responses in wheat

Summary Studies were made of days to ear emergence under the constant temperatures of 9, 14, 19 and 25°C and 16 h photoperiod in three sets of wheat lines each possessing genotypes differing for developmental responses.Days to ear emergence in three near-isogenic lines of the wheat cultivar Triple Dirk, which differed for vernalization response, increased as the strength of the response increased. At the four temperatures Triple Dirk D (Vrn 1 vrn 2) was not significantly different from normal Triple Dirk (Vrn 1 Vrn 2) but Triple Dirk B (vrn 1 Vrn 2) was significantly (P=0.01) later than normal Triple Dirk at each temperature. This indicates that the vrn 1 allele confers stronger vernalization response than vrn 2 over the range of temperatures (9–24°C). However, Triple Dirk C (vrn 1 vrn 2) failed to head after 120 days at each temperature indicating strong interaction between vrn 1 and vrn 2 with each other (and possibly the Triple Dirk back-ground) to give a much stronger vernalization response than predictions from additivity of their individual effects.The second set comprised the four Chinese Spring/Thatcher chromosome substitution lines CS/T 3B, 6B, 7B and 5D, plus Chinese Spring and Thatcher, and were grown in the unvernalized condition. CS/T 5D was similar in days to ear emergence as Chinese Spring at all four temperatures but the other three lines were earlier to ear emergence, particularly as the temperature increased. Days to ear emergence was fastest at 14°C in all lines, except CS/T 3B, in which it decreased progressively from 9 to 24°C.The third set of Chinese Spring and Thatcher and the homoeologous group 2 chromosomes of Thatcher substituted in Chinese Spring, the group which is considered to be involved in the control of photoperiod sensitivity. The three substitution lines responded differently to temperature compared with Chinese Spring and with each other, with chromosome 2D being the least, and chromosome 2B the most, responsive to temperature.     

The relationships of hardening period and the expression of frost resistance in chromosome substitution lines of wheat

Summary The highly frost resistant wheat variety Cheyenne (donor) and the poorly frost resistant variety Chinese Spring (recipient) were frozen at –9° C and –11° C at various stages of hardening, as were a number of substitution lines of these two varieties (CS/Ch 3A, CS/Ch 5A, CS/Ch 7A, CS/Ch 2B, CS/Ch 4B, CS/Ch 5B, CS/Ch 4D, CS/Ch 5D). Chromosomes 5A, 5B, 5D, 4B and 7A of Cheyenne increased the frost resistance of the recipient variety to varying extents. However, the frost resistance changed not only as a function of the different chromosomes, but also as a function of the duration of hardening, indicating that genes responsible for frost resistance are expressed differently during different phases of the hardening process.     

Crossability percentages of bread wheat landraces from Shaanxi and Henan provinces,China with rye

Summary The crossability percentages of 118 landraces of wheat (Triticum aestivum L.) from Shaanxi and Henan provinces, China with rye (Secale cereale L.) have been tested. 14 landraces possess a higher crossability percentage than Chinese Spring, 30 landraces have a similar to and 74 landraces have a lower crossability percentage than Chinese Spring or are non-crossable with rye. Most landraces with high crossability percentage occur in south part of Shaanxi, and the west and southwest of Henan, their geographical distribution in these regions fits up with that in the Sichuan province.     

Crossability percentages of bread wheat landraces from Sichuan Province,China with rye

Summary The crossabilities of 177 landraces of wheat (Triticum aestivum L.) from Sichuan Basin and its adjacent mountain ranges with rye (Secale cereale L.) have been tested. 16 landraces possess a higher crossability than Chinese Spring, 34 landraces have a similar and 127 landraces have a lower crossability than Chinese Spring or are non-crossable with rye. Most landraces with high crossability occur in Qinling Mountain and Dabashan Mountain Ranges in north of Sichuan and the valleys of Minjiang River, Fujiang River and Jialinjiang River in Sichuan Basin.     

Crossability percentages of bread wheat collections from Tibet,China with rye

Summary The crossability percentages of 282 accessions of wheat (Triticum aestivum L.) collected in Tibet, China with rye (Secale cereale L.) have been tested. Five collections have a similar to and 277 accessions have a lower crossability percentage than Chinese Spring or are non-crossable with rye. The accessions with high crossability percentage occur along the highway near Lhasa. No landraces with higher crossability than Chinese Spring and rare landraces with similar crossability to Chinese Spring indicated that the landraces in Tibet region are different from those in Sichuan, Shaanxi and Henan provinces in the distribution frequency of high crossability, and there is no distribution of recessive kr4 alleles.     

Factors affecting haploid production in wheat using the Hordeum bulbosum system. 3. Post-fertilization effects on embryo survival

Summary An investigation was made of factors affecting the frequency of development of fertilized ovules following wheat x Hordeum bulbosum hybridizations. Seed survival, a term used here to represent the relationship between seed set and fertilization, was shown to vary between the wheat genotypes tested, but was unrelated to the crossability genes present within the wheat genotype. A high seed survival was obtained in crosses involving the variety TH3929 and the single chromosome substitution line, Chinese Spring (Hope 5B). Chinese Spring showed a poor seed survival. Some influence of the H. bulbosum genotype on seed survival was detected. The two environmental factors studied, namely the post-pollination application of gibberellic acid and the ambient temperature, were both shown to affect seed survival. The application of gibberellic acid immediately after pollination or 1 and 2 days after pollination improved seed survival, although the extent of the response was dependent upon the number and timing of the application(s). A genotype-dependent response to the ambient temperature was observed, TH3929 showing a slight but insignificant reduction and Chinese Spring a significant improvement in seed survival with an increase in temperature from 20°C to 26°C. The physiological processes which may be influenced by GA and the ambient temperature and may thereby affect seed survival are discussed.     

Effect of alien 5R(5A) chromosome substitution on ear-emergence time and winter hardiness in wheat-rye substitution lines

Ear emergence time and response to vernalization were investigated in 12 alien substitution lines in which a pair of chromosomes 5A of recipient spring wheat cultivars was replaced by a pair of chromosomes 5R of Siberian spring rye ‘Onokhoiskaya’. The recipients were 12 spring cultivars of common wheat, each carrying different Vrn genes. Spring rye ‘Onokhoiskaya’ had the Sp1 (now called Vrn-R1) gene for spring growth habit located on chromosome 5R, but its expression was weaker. The Vrn-R1 gene had no effect on growth habit, ear emergence time and response to vernalization in wheat-rye substitution lines. Ears emerged significantly later in the 5R(5A) alien substitution lines than in the recipient wheat cultivars with the Vrn-A1/Vrn-B1/vrn-D1 or Vrn-A1/vrn-B1/Vrn-D1 genotypes. No difference in ear emergence time was found between most of the 5R(5A) alien substitution lines and the cultivars carrying the recessive vrn-A1 gene. The presence of the Vrn2a and Vrn2b alleles at the Vrn2 (now called Vrn-B1) locus located on wheat chromosome 5B was confirmed.The replacement of chromosome 5A by chromosome 5R in wheat cultivars ‘Rang’ and ‘Mironovskaya Krupnozernaya’, which carries the single dominant gene Vrn-A1, converted them to winter growth habit. In field studies near Novosibirsk the winter hardiness of 5R(5A) wheat–rye substitution lines of ‘Rang’ and ‘Mironovskaya Krupnozernaya’ was increased by 20–47% and 27–34%, respectively, over the recurrent parents.     

Wheat pollen dispersal under semiarid field conditions: potential outcrossing with Triticum aestivum and Triticum turgidum

Isolation distance is the main barrier to crop-to-crop gene-flow. A 3-year study assessed the maximum potential outcrossing under field conditions between two wheat cultivars (Triticum aestivum L.) and between wheat and durum wheat (Triticum turgidum L. var. durum). Outcrossing was measured by seed set on emasculated recipient plants placed at four sides with different distances from a 3 m × 3 m T. aestivum (cultivar Chinese Spring) pollen source. Frequencies of seed set at 0 m distance were 45% (37–56%) for T. aestivum cultivars and 18% (5–30%) with T. turgidum. These values agree with hybridization in non-limiting pollen conditions measured by manual crosses in greenhouse. The number of pollen grains and the outcrossing frequencies decreased at increasing distances influenced by the prevailing wind direction. Under semiarid conditions of this assay, viable pollen was found 14 m from the pollen source, with a maximum distance of 8 m at which cross-pollination decreases below 1%. Ambient conditions affect pollen viability, hybridization and pollen dispersal. Data presented in this paper emphasize the major role played by environmental conditions in outcrossing. Data obtained in one area may therefore not coincide with the prevailing situation in different locations and climates.     

Chromosome locations of genes controlling ‘purple leaf base’ in rye and wheat

Summary Purple leaf base is expressed only if there is anthocyanin pigmentation in coleoptiles either in rye or in rye-wheat-additions. Genes controlling purple leaf base were found to be located on chromosomes 5R (An5), 4B (Ra2) and 6B (Ra3) using the trisomic set of rye cv. Esto and autoplasmic rye-wheat-additions, respectively.     

The effect of pollen load and pollen grain competition on fertilization success and progeny performance inFagopyrum esculentum

Summary Flowers of cultivated buckwheat (Fagopyrum esculentum Moench) often receive natural pollen loads of fewer than 10 pollen grains. The cultivated varieties also have high genetic variability. These observations raise the question of whether seed production in buckwheat is often limited by pollen delivery, and whether small increases in pollen load could result in gametophytic selection through pollen grain competition. In greenhouse-grown buckwheat plants, embryo sac penetration by pollen tubes was universal with 10 or more pollen grains. However, seed production increased with pollen load up to 30 grains per flower. Larger pollen loads, which intensify selection among gametophytes, resulted in more vigorous progeny. Seedlings produced from high pollen load (15–20 pollen grains) were larger (40% by weight) than those from low pollen load (5 pollen grains). These results are evidence that pollen grain competition can occur in buckwheat with benefits for progeny performance.     

Inter-varietal pollen tube growth and ovule penetration in the avocado

Summary Results are presented showing number of pollen tubes in the pistil, and ovule penetrations in crosses involving the avocado varieties Edranol, Ryan, Hass, Reed, Talbot, Jalna, Fuerte, Bacon and Sharwil. Between 5% and 100% of the ovules in each cross were penetrated by a pollen tube. The female parent exerted more control than the male on numbers of tubes and ovule penetrations and the results for type A varieties were more consistent than those for type B varieties. Penetration of an ovule by two tubes was observed in all varieties except Reed and Fuerte and penetration of an ovule by three tubes was observed in the variety Edranol only. With the possible exception of the variety Bacon as female parent there was no difference between pollination with pollen from type A or B varieties.     

Chromosome location of genes controlling the content of dietary fibre and arabinoxylans in rye

Differences in fruit-set success reputed to exist between Heveabrasiliensis genetic clones were verified by an analysis of breeding recordsfor the clones PB 5/51, RRIM 600 and PR 107. Although the bestfruit-set success was obtained with PB 5/51 following artificial (hand)pollination, this clonal trait was not reflected in terms of greater numbersof pollen tubes developing it the styles of hand-pollinated PB 5/51 femaleflowers. It was observed, however, that more pollen tubes reached theovules in PB 5/51 than in RRIM 600 or PR 107. Significantly, PB 5/51female flowers required fewer pollen tubes to effect penetration of all threeof its ovules. As fruit formation in Hevea is dependent on all threeovules of the flower being successfully fertilised, PB 5/51 female flowershave hence a greater propensity for successful fruit-set. In all the threeclones studied, the frequency distribution of female flowers with 0, 1, 2 or3 of the ovules penetrated by pollen tubes did not conform to binomialexpectations. Flowers with no ovule penetrated and flowers with all threeovules penetrated were greatly over-represented. One explanation for thisnon-random distribution is the existence of `receptive' female flowers thatfavour successful fertilisation whereas `non-receptive' flowers tend to remainunfertilised even when hand-pollinated.