Research Progress of Regulatory Factors for Tillering and Earring of Large-spike and Multispike Cultivars of Winter Wheat

Na’na LI, Yanyan PU, Yongchao GONG, Xinhua LI, Peng LI, Hanfeng DING*

1. Shandong Centre of Crop Germplasm Resources, Ji’nan 250100, China;

2. Institute of Atomic Energy Application in Agriculture, Shandong Academy of Agricultural Sciences, Ji’nan 250100, China

Wheat (Triticum aestivum L.)is the most important food crop worldwide, its high and stable yield plays an important role for ensuring efficient supply of grain, social stability and economic development. The yield structure of wheat is composed of three elements: spike number, kernels per spike and 1 000-kernel weight, and is a quantitative character controlled by multiple genes,its phenotypic characteristic is easily affected by environment[1-2], therefore,its genetic improvement is restricted by many factors. Based on years of investigation and study in different areas, the predecessors[3]put forward that spike number was the dominant factor for yield composition. So increasing effective spike number was the precondition of improving yield per unit area. Tillering, growth and earring played an important regulating role in group structure and yield,thus improving tillering quantity and quality can increase ear number obviously.

In recent years, the application of wheat varieties with different genotypes in super high-yield field has always been the hot topic of related research[4-6].At the present stage,superhigh-yielding breeding of wheat has been mostly dependent on multi-spike cultivars. But with the release of fine varieties and the improvement of cultivation measures, the yield of largespike cultivars can also reach 9 000 kg/hm2. Therefore, the application trend of different cultivars may change from pure middle-spike and multi-spike cultivars to middle-spike, multi-spike and large-spike cultivars[7]. But largepike cultivars have low rates of tiller and earbearing tiller, which is an important factor restricting yield, thereby extremely limiting the play of produc-tive potential. How to improve tillering and earring rate, and ensure enough spike number per unit area, is the problem to be solved[8].Tiller is an important biological characteristic of wheat, and an important element determining colonies and quality of wheat in high-yield cultivation[9], as well as related to group internal farmland microclimate and the construction of rational colony structure[10-13], so we can promote and control tiller growth to regulate rational colony structure in production, thus exploring ways for high-yielding[14-15]. For years,people have studied and analyzed the dynamic variation rule of tillering and earring of winter wheat. Synthesizing existing research results, the paper summarized the characteristics of tillering and earring of winter wheat with different spike cultivars,to provide theoretical basis and technical index for colony regulation and control of high-yield cultivation of different cultivars.

Effect of Genetic Factors on the Characteristics of Tillering and Earring of Winter Wheat

Tillering character is a quantitative character controlled by multiple genes,genetic factor has a particularly important effect on tillering. The hereditary characters of the cultivars determine spike number[16-18]. Different wheat cultivars with different genotypes have different tillering abilities, earbearing tiller rates and spike number.The predecessors did many researches on genetic factors of wheat tillering[19],and found the genes which inhibited tillering, such as tin[20], tin-1[21], tin2[22],tin3[23].Li et al.[24]found that there were related genes in a region near Gli2A2(Xp sr10)in the short arm of chromosome 6A,which seriously affected tiller number of wheat and seedling growth.Zhang Qianhui et al.[25-26]showed that effective tillering number of wheat had high main gene heritability and modified by multiple genes,but mainly controlled by two major genes, and the two genes had interaction effect, one of the two can inhibit effective tillering.For main control factors affecting tillering and earring, Mi Guohua et al.[27-28]had different opinions,they considered that hereditary characters of wheat cultivars determined the effective tillering number of wheat. Wang Shaozhong et al. proved that low percentage of earbearing tiller was the hereditary character of parts of largespike wheat cultivars, can not be changed easily by cultivation measures.

Effect of Group Environment Factors on the Characteristics of Tillering and Earring of Winter Wheat

The former research results[29-33]indicated that the characters of tillering and earring of different cultivars were affected greatly by group environment factors, but different cultivars had different response sensitivities to environment, therefore, in actual production, we can regulate and control according to different response sensitivities of different cultivars to environment,to ensure the formation of rational group structure of each cultivar,thus achieving high yield. For some large-spike cultivars,the change of environmental factors and cultivation means can bring noticeable change to ear number; comparing with genetic factors, group environment had a greater impact on planting distance,sowing time, seeding density and endogenous hormones[7,32].

Row-spacing

After determining basic seedlings,rational row-spacing will have a significant positive effect on yield. When the group yield reaches the highest, the row-spacing in the field will be the best.Rowing-spacing,as an important means of regulation and control, has always received special attention from cultivation experts[34-36].The traditional seeding way was sowing in drill,different row-spacing generated different characteristics of tillering and earring of different cultivars. Wu Yu’e et al.[37]suggested that to achieve the maximum number of spike, different cultivars must have different optimum spacing, specifically, the optimum spacing for large-spike cultivars was 16.7 cm, and the optimum spacing for compact and loose multi-spike cultivars was 20 and 23.3 cm,respectively.Sun Hongyong et al.[38]agreed that spike number of winter wheat decreased with the increase of row spacing; the narrower the row spacing, the more the spike number. Therefore,narrowing the row spacing can make wheat plane distribution uniform and effectively inhibit ineffective tillering for better using surrounding resources,but the increase potential of final grain yield was closely related to cultivars and environment.

At the same time, some scholars put forward some corresponding planting modes[39-40]according to the characteristics of different ecological regions. Wang Fahong et al.[41]proved that comparing with convention planting, raised bed planting for wheat was conducive to taking full advantage of border row,although spike number per unit area showed a slight decrease,kernels per spike and 1 000-kernel weight were improved prominently,which optimized wheat yield structure and achieved yield increase. Liu Yinjie et al.[42]found that wide-narrow row planting of wheat can give full play to the border row,and realize the harmonious development of low group,strong individual and yield increase,which was beneficial to lodging-resistance, premature senescence resistance and yield increase.

Sowing date

Appropriate sowing time, as an important cultivation technique, not only ensures the synchronization of normal and effective production and growth progress with the optimum season, but also helps coordinate the relationship between growing development of crops and environmental conditions, makes full use of natural resources and avoids adverse weather[43-45].Sowing date is the primary means for adjusting wheat tillering and effective accumulative temperature, and is very important for the formation of spikes.

Judging the most suitable sowing date of wheat should be based on accumulated temperature last year,variety properties and soil moisture.In the meantime, there are a lot of differences in different cultivars,such as the adaptability to ecological environment,and the requirements for appropriate temperature, light and water condi-tions.Thus,we should select the most suitable varieties according to different ecological environment, to match different sowing dates, thus making full use of local temperature, light, water and thermal resources, finally achieving yield and income increase. Early sowing is conducive to making seedlings strong before winter, promoting root and overground part development, and improving tiller number; moreover, it is also good for increasing dry matter accumulation and improving grain filling rate. For winter wheat, if it is less affected by cold damage during wintering period,winter wheat can turn green early, and has the advantages of long growth time of seedling stage and more accumulated temperature, the growth and development of winter wheat can be improved,thereby increasing spike number.If the sowing date is late, the growth of seedlings will be slow before winter,thereby generating insufficient tillers and less groups[46-47].Han Jinling et al.[48]demonstrated that along with the delay of planting date, spikelet number per stem and fertile spikelets decreased significantly,thus effective spike number decreased markedly. But we can supplement basic seedlings to make up for the insufficient of accumulated temperature before winter, thereby increasing spike number per unit area.Zhang Xuepin et al.[49]showed that total tiller number decreased obviously with the delay of sowing date, but the percentage of earbearing tiller improved conversely. The effect of the treatment during sowing time and tillering stage was far greater than genotype difference, the formation of tillering was mainly dependent on the control of ecological factors.

Planting density

To achieve high yield, both individual and group qualities should be improved. Planting density affects the group structure of wheat, thereby causing the differences of field climatic conditions, such as temperature, light and humidity, etc., and then influencing crops individual and group qualities from the aspect of vegetative growth,thereby determining the group development trend of whole growth period to a certain degree. Therefore, planting density is the most basic measure for adjusting group and individual development[50-53].Planting density has an obvious effect on all growth indicators and aboveground dry weight during wheat tillering stage, too high density will cause slow growth,which is mainly reflected in the decrease of the number of blades and root system,thin and weak seedlings, etc.[54]. In a certain range, multi-spike is beneficial to yield increase, but if out of the range, average spike weight does not rise but decreases. Chai Shouxi et al.[55]studied the influence of planting density on winter wheat yield and related indicators, and found that 390 plants/m2planting density can reach the highest grain yield. To establish rational group structure, we should try to reduce basic seedlings under the premise of enough spikes, make individual develop fully and improve the percentage of earbearing tiller, thereby coordinating internal conflicts within group and achieving high yield[56-57].

Wang Zhijie et al.[58]researched the characteristics of tillering and earring of multi-spike cultivar (Yumai 49)of wheat, the results indicated that there was a significant difference in spike number under different density treatments,and spike number showed an increasing trend with the increase of basic seedlings, however, kernels per spike and 1 000-grain weight had a reducing trend with the increase of basic seedlings. Wang Longjun et al.[59]argued that along with the increase of density, spike number per 667 m2did not necessarily increase,in certain environment conditions, spike number per unit area of specific varieties had a relative stable upper limit. Zhu Yunji et al.[29]proved that the rate of tillering and earring of large-spike cultivar(Lankao 906)of wheat was low, increasing density can increase spike number, but would generate the decrease of kernels per spike and the reduction of kernel weight as well as low yield. Thus, in the production practice,we should control planting density according to different cultivars; for multispike cultivars, the relationship between group and individual should be coordinated by controlling suitable group, to reach the effects of the increase of kernels per spike and the improvement of kernel weight, finally,the coordinated development of the three elements was achieved; largespike cultivars had more kernels per spike, but the low percentage of earbearing tiller was easy to cause insufficient spikes, thereby affecting wheat yield; so appropriately increasing seeding rate can effectively increase spike number,thus improving yield.

Effect of Endogenous Hormones on the Characteristics of Tillering and Earring of Winter Wheat

With the deepening of the research, the relationship of tillering and earring with endogenous hormone has become the focus of many scholars[60].Endogenous hormone plays an important role in tillering and its growth[61].The reasons for tiller senescence are the competition of assimilated products and hormone or the change of hormonal equilibrium. External environment can control the characteristics of tillering and earring by changing hormone content and equilibrium in plants[62]. Therefore,the hormonal equilibrium in plants is the principal element regulating plant growth[63-66].

The polarization of wheat tillering has a close relationship with hormone levels, plant hormones have an apparent regulatory effect on tillering growth[67]. Li Chunxi et al.[68-70]argued that during the tillering process of different winter wheat cultivars, the dynamic change of the contents of endogenous indoleacetic acid (IAA),zeatin riboside(ZR)and zeatin(Z)was obvious. So the dynamic change of wheat tillering was closely related to plant endogenous hormones IAA and(ZR+Z) contents, especially IAA/(ZR+Z)value and its variation trend. In the initial stage of tillering, tillering ability had a significant negative correlation with IAA/(ZR+Z),the lower or declining IAA/(ZR+Z)value was more favorable to tillering, conversely, the higher or rising ratio may inhibit tillering or lead to tiller senescence. Zhu Yunji et al.[60]showed that there were apparent differences in endogenous hormone contents of tillering and stem for different spikes cultivars. At peak tillering stage,endogenous hormones IAA and(ZR+Z)contents in stem of large-spikecultivar Yumai 66 were less than these of multi-spike cultivar Yumai 49, however,for abscisic acid (ABA), the former was higher than the latter,and the gap between stem and tillering was larger than that of Yumai 49, thus the stem of Yumai 66 was clearly better than tillering, and tiller number of Yumai 66 was less than that of Yumai 49;at the tiller senescence stage, the change of endogenous hormone(ZR+Z) of large-spike cultivar Yumai 66Ⅰand Ⅱ happened at double ridge stage, then the ear differentiation began to slow down, and the gap with stem leaf age enlarged; ABA change before shape senescence aggravated the process; however, the change of endogenous hormones in tillering and stem of multi-spike cultivar Yumai 49ⅠandⅡwas synchronous, so the gap value was stable, finally forming spikes.

Problems and Prospect

As the measure standard of individual strong degree, the quantity and quality of tillering and earring of wheat had a close relationship with the rationality of group dynamic development.Although many researches have focused on tillering and earring characters of different spike cultivars of winter wheat,there were still some aspects to be further discussed, thus we should analyze the relationship between tillering and influencing factors from multiangles, grasp the occurrence rule of tillering, thereby improving the rate of tillering and earring, increasing spike number per unit area,achieving the increase from both production and income of different spike cultivars of winter wheat.

Firstly, the tillering formation of different spike cultivars of winter wheat was a very complicated process, and affected by such factors as temperature, illumination, nutrition, heredity and hormone, etc. The present research on tillering has been focused only on the effect of single element on single cultivar, but if we want to effectively control tillering, we must make clear the occurrence mechanism of tillering and the relationship among different factors, then in-depth study the occurrence of tillering,only in this way,can we control it better.

Secondly, the former researches on tillering and earring characters of different spike cultivars of winter wheat were mostly concentrated on such environmental factors as sowing date,planting density and row-spacing,etc.,however, the studies on endogenous hormone content, hormonal equilibrium as well as exogenous hormone application during the tillering process were fewer.So to strengthen the study on these aspects in the future will help us gain a better understanding of tillering occurrence mechanism.

Thirdly,the reports on the study of the relationship of genetic expression and control with wheat tillering,and the control of tillering using molecule means were fewer. From now on , if we can make progress in studying the relationship between molecular level and wheat tillering, it will help us indepth understand the tillering occurrence mechanism.

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