Formation Of Abscission Layer Research Articles

Response of Abscission Zone of Blue Honeysuckle (Lonicera caerulea L.) Fruit to GA3 and 2,4-D Spray Application

The nutritional value of blue honeysuckle (Lonicera caerulea L.) fruit is abundant; however, its production faces challenges due to a short harvesting period and fruit drop issues. In this study, the effects and potential mechanisms of two different plant growth regulators, GA3 (Gibberellins acid) and 2,4-D (2,4-Dichlorophenoxyacetic acid), on blue honeysuckle fruit abscission and abscission layer formation were investigated. The main cultivated variety of blue honeysuckle, ‘Berel’, was used as the experimental material. GA3 and 2,4-D were sprayed on the plants from the veraison. The anatomical structure of the fruit abscission zone (FAZ) was observed after treatment. Key enzymes involved in cell wall degradation, pectinase, cellulase, polygalacturonase, and pectin methylesterase, were analyzed for their activities. Furthermore, the gene expression levels of cell wall detachment-related genes CX1, CX2, PL20, PE, and key genes for gibberellin and ethylene synthesis GA2OX1, GA1, ACO, and ACO3 were examined. The results indicated that the application of GA3 and 2,4-D could delay the formation of the abscission layer. In the FAZ treated with GA3 and 2,4-D at 5 DAT, the activity of key enzymes involved in cell wall degradation decreased, the expression of genes related to cell wall degradation enzymes and key genes for ethylene synthesis was inhibited, and the drop of fruit reduced. In conclusion, exogenous application of GA3 and 2,4-D suppresses the abscission of ‘Berel’ blue honeysuckle fruit, likely through the inhibition of cell wall degradation and abscission layer formation.

A Novel Introgression Line Library Derived from a Wild Melon Gives Insights into the Genetics of Melon Domestication, Uncovering New Genetic Variability Useful for Breeding.

A collection of 30 melon introgression lines (ILs) was developed from the wild accession Ames 24297 (TRI) into 'Piel de Sapo' (PS) genetic background. Each IL carried an average of 1.4 introgressions from TRI, and the introgressions represented 91.4% of the TRI genome. Twenty-two ILs, representing 75% of the TRI genome, were evaluated in greenhouse (Algarrobo and Meliana) and field (Alcàsser) trials, mainly to study traits related to domestication syndrome such as fruit weight (FW) and flesh content (FFP), as well as other fruit quality traits as fruit shape (FS), flesh firmness (FF), soluble solid concentration (SSC), rind color and abscission layer. The IL collection showed an impressive variation in size-related traits, with FW ranging from 800 to 4100 g, reflecting the strong effect of the wild genome on these traits. Most of the ILs produced smaller fruits compared with PS; however, unexpectedly, the IL TRI05-2 produced bigger fruits, likely due to new epistatic interacions with the PS genetic background. In contrast, the genotypic effect for FS was smaller, and few QTLs with notable effects were detected. Interestingly, variability was also observed for FFP, FF and SSC, rind color and abscission layer formation. Genes in these introgressions are candidates for having been involved in melon domestication and diversification as well. These results confirm that the TRI IL collection is a very powerful tool for mapping traits of agronomic interest in melon, allowing the confirmation of previously reported QTLs and the identification of new ones to better understand the domestication process of this crop.

A stepwise route to domesticate rice by controlling seed shattering and panicle shape

Asian rice (Oryza sativa L.) is consumed by more than half of the world's population. Despite its global importance, the process of early rice domestication remains unclear. During domestication, wild rice (Oryza rufipogon Griff.) acquired non-seed-shattering behavior, allowing humans to increase grain yield. Previous studies argued that a reduction in seed shattering triggered by the sh4 mutation led to increased yield during rice domestication, but our experiments using wild introgression lines show that the domesticated sh4 allele alone is insufficient for shattering loss in O. rufipogon. The interruption of abscission layer formation requires both sh4 and qSH3 mutations, demonstrating that the selection of shattering loss in wild rice was not as simple as previously suggested. Here we identified a causal single-nucleotide polymorphism at qSH3 within the seed-shattering gene OsSh1, which is conserved in indica and japonica subspecies but absent in the circum-aus group of rice. Through harvest experiments, we further demonstrated that seed shattering alone did not significantly impact yield; rather, yield increases were observed with closed panicle formation controlled by SPR3 and further augmented by nonshattering, conferred by integration of sh4 and qSH3 alleles. Complementary manipulation of panicle shape and seed shattering results in a mechanically stable panicle structure. We propose a stepwise route for the earliest phase of rice domestication, wherein selection of visible SPR3-controlled closed panicle morphology was instrumental in the sequential recruitment of sh4 and qSH3, which together led to the loss of shattering.

Chromosome-level genome assembly of Zizania latifolia provides insights into its seed shattering and phytocassane biosynthesis

Chinese wild rice (Zizania latifolia; family: Gramineae) is a valuable medicinal homologous grain in East and Southeast Asia. Here, using Nanopore sequencing and Hi-C scaffolding, we generated a 547.38 Mb chromosome-level genome assembly comprising 332 contigs and 164 scaffolds (contig N50 = 4.48 Mb; scaffold N50 = 32.79 Mb). The genome harbors 38,852 genes, with 52.89% of the genome comprising repetitive sequences. Phylogenetic analyses revealed close relation of Z. latifolia to Leersia perrieri and Oryza species, with a divergence time of 19.7–31.0 million years. Collinearity and transcriptome analyses revealed candidate genes related to seed shattering, providing basic information on abscission layer formation and degradation in Z. latifolia. Moreover, two genomic blocks in the Z. latifolia genome showed good synteny with the rice phytocassane biosynthetic gene cluster. The updated genome will support future studies on the genetic improvement of Chinese wild rice and comparative analyses between Z. latifolia and other plants.

Studies of Vegetative Growth, Inflorescence Development and Eco-Dormancy Formation of Abscission Layers in Streptocarpus formosus (Gesneriaceae)

Streptocarpus formosus (Hilliard & B.L. Burtt) T.J. Edwards is a flowering herbaceous perennial indigenous to South Africa and is part of the rosulate group of herbaceous acaulescent plants within the Gesneriaceae family. According to the National Assessment database for the Red List of South African Plants version 2020.1., the plant is listed as rare. The ornamental use of S. formosus has untapped commercial potential as a flowering indoor pot plant, an outdoor bedding plant for shade and as a cut flower for the vase, all of which are limited by a five-month eco-dormancy period during the late autumn and all through the cold season in the short-day winter months. Viable commercial production will require cultivation techniques that produce flowering plants all year round. This study investigated the effectiveness of applying root zone heating to S. formosus plants grown in deep water culture hydroponics during the eco-dormancy period in preventing abscission layer formation and in encouraging flowering and assessed the growth activity response of the plants. The experiment was conducted over eight weeks during the winter season in the greenhouse at Kirstenbosch Botanical garden in water reservoirs, each maintained at five different experimental temperature treatments (18, 22, 26—control, 30 and 34 °C) applied to 10 sample replicates. The results showed that the lowest hydroponic root zone temperature of 18 °C had the greatest effect on the vegetative growth of S. formosus, with the highest average increases in fresh weight (1078 g), root length (211 cm), overall leaf length (362 cm) and the number of newly leaves formed (177 = n), all noted as statistically significant when compared with the other water temperature treatments, which yielded negative results from reduced vegetative growth. Findings from the study also revealed that while all heated solutions significantly prevented the formation of abscission layers of S. formosus, they had a less significant effect on inflorescence formation, with only 18 °C having the greatest positive effect on flower development.

Same Locus for Non-shattering Seed Pod in Two Independently Domesticated Legumes, Vigna angularis and Vigna unguiculata.

Loss of pod shattering is one of the most important domestication-related traits in legume crops. The non-shattering phenotypes have been achieved either by disturbed formation of abscission layer between the valves, or by loss of helical tension in sclerenchyma of endocarp, that split open the pods to disperse the seeds. During domestication, azuki bean (Vigna angularis) and yard-long bean (Vigna unguiculata cv-gr. Sesquipedalis) have reduced or lost the sclerenchyma and thus the shattering behavior of seed pods. Here we performed fine-mapping with backcrossed populations and narrowed the candidate genomic region down to 4 kbp in azuki bean and 13 kbp in yard-long bean. Among the genes located in these regions, we found MYB26 genes encoded truncated proteins in azuki bean, yard-long bean, and even cowpea. As such, our findings indicate that independent domestication on the two legumes has selected the same locus for the same traits. We also argue that MYB26 could be a target gene for improving shattering phenotype in other legumes, such as soybean.

Detection of a novel locus involved in non-seed-shattering behaviour of Japonica rice cultivar, Oryzasativa 'Nipponbare'.

A novel locus, qCSS3, involved in the non-seed-shattering behaviour of Japonica rice cultivar, 'Nipponbare', was detected by QTL-seq analysis using the segregating population with the fixed known seed-shattering loci. Asian cultivated rice, Oryzasativa, was domesticated from its wild ancestor, O.rufipogon. Loss of seed shattering is one of the most recognisable traits selected during rice domestication. Three quantitative trait loci (QTLs), qSH1, qSH3, and sh4, were previously reported to be involved in the loss of seed shattering of Japonica cultivated rice, O.sativa 'Nipponbare'. However, the introgression line (IL) carrying 'Nipponbare' alleles at these three loci in the genetic background of wild rice, O.rufipogon W630, showed a lower value for detaching a grain from the pedicel than 'Nipponbare'. Here, we investigated abscission layer formation in the IL and found a partially formed abscission layer in the central region between the epidermis and vascular bundles. Based on QTL-seq analysis using the F2 population obtained from a cross between 'Nipponbare' and the IL, we detected two novel loci qCSS3 and qCSS9 (QTL for the Control of Seed Shattering in rice on chromosomes 3 and 9), which were found to be involved in the difference in seed-shattering degree between 'Nipponbare' and W630. Then, we further focused on qCSS3 in order to understand its potential role on the loss of seed shattering. The candidate region of qCSS3 was found to be located within a 526-kb region using substitution mapping analysis. Interestingly, the qCSS3 candidate region partially overlaps the selective sweep detected for Japonica but not for Indica rice cultivars, suggesting that this region harbours the mutation at a novel seed-shattering locus specifically selected for non-seed-shattering behaviour in Japonica cultivars.

Synchrony in fall leaf drop: chlorophyll degradation, color change, and abscission layer formation in three temperate deciduous tree species

Deciduous tree species remove some nutrients from their leaves during fall leaf senescence through retranslocation. Retranslocation impacts the timeline of leaf fall, amount of active chlorophyll, and overall leaf nitrogen content as fall color change occurs. Our objective was to identify interspecific differences in the timing of abscission layer formation, leaf color change, and the level of chlorophyll degradation of young trees during fall senescence. Leaf relative chlorophyll content for three tree species was measured during fall 2015 by a greenness meter. These measurements were calibrated for each species through spectrophotometric determination of leaf chlorophyll concentration. Abscission layer formation was tracked using light microscopy of sampled leaves. Excised leaves were photographed on a flat white surface to track species leaf color through time. All three species had different chlorophyll declination rates throughout the fall season. The maple species started with less chlorophyll and began abscission layer formation earlier. The other two species had a similar starting chlorophyll level and onset timing of abscission layer formation. Visible leaf color change was not associated with a threshold in either chlorophyll degradation or abscission layer formation across species. Maple species degraded less chlorophyll on average, in the fall, than did the oak and beech species. The rate of chlorophyll degradation in coordination with abscission layer formation varied by species. Color change was not a good predictor of level of chlorophyll degradation in leaves across species.

Identification and expression of genes associated with the abscission layer controlling seed shattering in Lolium perenne.

Perennial ryegrass (Lolium perenne) is one of the most important pasture grasses in the world. However, seed production is negatively impacted by the seed shattering (shedding) nature of this species. Recently, genes involved in the seed shattering process have been isolated and functionally characterized in several crop species. The aim of this study was to identify the genes playing critical roles in the seed shattering process in perennial ryegrass. DNA sequences of genes involved in seed shattering in the Poaceae were used to identify and isolate target genes in perennial ryegrass using a comparative genomics strategy. The candidate seed shattering genes were identified using an ‘in-house’ perennial ryegrass transcriptome database. The relative expression levels of the candidate ryegrass shattering genes were determined using RT–qPCR during different floret and seed developmental stages. Histological analysis of the abscission layer was also conducted. Homologues of seed shattering genes were identified and isolated from perennial ryegrass, and the relative gene expression results suggested that several genes, including LpqSH1 and LpSH1, might have a role in abscission layer formation during seed development. In addition, lignification of the abscission layer may play an important role in the abscission process. A genetic model for seed shattering in perennial ryegrass is suggested and may be useful for directing gene editing towards the production of a reduced-shattering ryegrass.

Estimation of loci involved in non-shattering of seeds in early rice domestication.

Rice (Oryza sativa L.) is widely cultivated around the world and is known to be domesticated from its wild form, O. rufipogon. A loss of seed shattering is one of the most obvious phenotypic changes selected for during rice domestication. Previously, three seed-shattering loci, qSH1, sh4, and qSH3 were reported to be involved in non-shattering of seeds of Japonica-type cultivated rice, O. sativa cv. Nipponbare. In this study, we focused on non-shattering characteristics of O. sativa Indica cv. IR36 having functional allele at qSH1. We produced backcross recombinant inbred lines having chromosomal segments from IR36 in the genetic background of wild rice, O. rufipogon W630. Histological and quantitative trait loci analyses of abscission layer formation were conducted. In the analysis of quantitative trait loci, a strong peak was observed close to sh4. We, nevertheless, found that some lines showed complete abscission layer formation despite carrying the IR36 allele at sh4, implying that non-shattering of seeds of IR36 could be regulated by the combination of mutations at sh4 and other seed-shattering loci. We also genotyped qSH3, a recently identified seed-shattering locus. Lines that have the IR36 alleles at sh4 and qSH3 showed inhibition of abscission layer formation but the degree of seed shattering was different from that of IR36. On the basis of these results, we estimated that non-shattering of seeds in early rice domestication involved mutations in at least three loci, and these genetic materials produced in this study may help to identify novel seed-shattering loci.

Mapping of Novel QTL Regulating Grain Shattering Using Doubled Haploid Population in Rice (Oryza sativa L.).

The critical evolutionary step during domestication of major cereals was elimination of seed shattering because the easy-to-shatter trait in wild relatives results in a severe reduction in yield. In this study, we analyzed the QTLs associated with shattering employing a high-density genetic map in doubled haploid (DH) population of rice (Oryza sativa L.). A genetic linkage map was generated with 217 microsatellite markers spanning 2082.4 cM and covering 12 rice chromosomes with an average interval of 9.6 cM between markers based on 120 DHLs derived from a cross between Cheongcheong indica type cultivar and Nagdong japonica type cultivar. In the QTL analysis, five QTLs pertaining to the breaking tensile strength (BTS) were detected in 2013 and 2015. Two regions of the QTLs related to BTS on chromosome 1 and chromosome 6 were detected. Several important genes are distributed in 1 Mbp region of the QTL on chromosome 6 and they are related to the formation of abscission layer. We decide to name this QTL qSh6 and the candidate genes in the qSh6 region can be employed usefully in further research for cloning.

Inhibition of abscission layer formation by an interaction of two seed-shattering loci, <i>sh4</i> and <i>qSH3</i>, in rice

Loss of seed shattering was one of the key phenotypic changes selected for in the domestication of many crop species. Asian cultivated rice, Oryza sativa L., was domesticated from its wild ancestor, O. rufipogon, and three seed-shattering loci, qSH1, sh4 and qSH3, have been reported to be involved in the loss of seed shattering in cultivated rice. Here, we analysed the seed-shattering behaviour of wild rice using introgression lines carrying the cultivated alleles from O. sativa Nipponbare in the genetic background of wild rice, O. rufipogon W630. We first carried out fine mapping of the qSH3 region and found that the qSH3 locus is localized in an 850-kb region on chromosome 3. We then analysed the effects of the Nipponbare alleles at sh4 and qSH3 on seed-shattering behaviour in wild rice, as a mutation at qSH1 was not commonly found in rice cultivars. Seed-shattering behaviour did not change in the two types of introgression line independently carrying the Nipponbare-homozygous alleles at sh4 or qSH3 in the genetic background of wild rice. However, the introgression lines having the Nipponbare-homozygous alleles at both sh4 and qSH3 showed a reduction in the degree of seed shattering. Histological and scanning electron microscopy analyses revealed that abscission layer formation was inhibited around the vascular bundles in these lines. Since the qSH3 region, as well as sh4, has been shown to be under artificial selection, the interaction of mutations at these two loci may have played a role in the initial loss of seed shattering during rice domestication.

Effect of quantitative trait loci for seed shattering on abscission layer formation in Asian wild rice Oryza rufipogon.

Asian cultivated rice Oryza sativa L. was domesticated from its wild ancestor, O. rufipogon. During domestication, the cultivated rice lost its seed-shattering behaviour. Previous studies have shown that two major quantitative trait loci (QTLs; qSH1 and sh4) are responsible for the seed-shattering degree. Here, we produced introgression lines carrying non-functional alleles from O. sativa ‘Nipponbare’ at the two major QTLs in the genetic background of wild rice O. rufipogon W630, and examined the effects of the two QTLs on seed shattering and abscission layer formation. The introgression lines, with Nipponbare alleles at either or both loci, showed complete or partial abscission layer formation, respectively, indicating that other unknown loci might be involved in enhancing seed shattering in wild rice. We detected a single QTL named qSH3 regulating seed-shattering degree using an F2 population between Nipponbare and the introgression line carrying Nipponbare alleles at the two QTLs. Although we generated an introgression line for qSH3 alone, no effects on seed shattering were observed. However, a significant effect on seed-shattering degree was observed for the introgression line carrying Nipponbare alleles at qSH3 and the two QTLs, suggesting an important role of qSH3 on seed shattering in coordination with the two QTLs.

Timing is everything: early degradation of abscission layer is associated with increased seed shattering in U.S. weedy rice

BackgroundSeed shattering, or shedding, is an important fitness trait for wild and weedy grasses. U.S. weedy rice (Oryza sativa) is a highly shattering weed, thought to have evolved from non-shattering cultivated ancestors. All U.S. weedy rice individuals examined to date contain a mutation in the sh4 locus associated with loss of shattering during rice domestication. Weedy individuals also share the shattering trait with wild rice, but not the ancestral shattering mutation at sh4; thus, how weedy rice reacquired the shattering phenotype is unknown. To establish the morphological basis of the parallel evolution of seed shattering in weedy rice and wild, we examined the abscission layer at the flower-pedicel junction in weedy individuals in comparison with wild and cultivated relatives.ResultsConsistent with previous work, shattering wild rice individuals possess clear, defined abscission layers at flowering, whereas non-shattering cultivated rice individuals do not. Shattering weedy rice from two separately evolved populations in the U.S. (SH and BHA) show patterns of abscission layer formation and degradation distinct from wild rice. Prior to flowering, the abscission layer has formed in all weedy individuals and by flowering it is already degrading. In contrast, wild O. rufipogon abscission layers have been shown not to degrade until after flowering has occurred.ConclusionsSeed shattering in weedy rice involves the formation and degradation of an abscission layer in the flower-pedicel junction, as in wild Oryza, but is a developmentally different process from shattering in wild rice. Weedy rice abscission layers appear to break down earlier than wild abscission layers. The timing of weedy abscission layer degradation suggests that unidentified regulatory genes may play a critical role in the reacquisition of shattering in weedy rice, and sheds light on the morphological basis of parallel evolution for shattering in weedy and wild rice.

PFA CONTROL IN 'SERR' IN CHILE

Walnut varieties to various degrees present pistillate flowers abortion (PFA), due to an excess of pollen that generates ethylene, causing abscission layer formation and organ excision. 'Serr' is one of the most affected walnut varieties by this disorder. In Chile, PFA seasonal records have registered up to 78% incidence in some growing areas. From the 2004-2005 season up till the present, Aminoethoxyvinylglycine (AVG) had been evaluated for 'Serr' PFA control, in order to determine wetting requirements, chemical dosages, and treatment timings. A dose of 125 grams a.i. per hectare resulted in good PFA control. Half of that concentration showed erratic performance, and a double dose did not improve PFA control significatively. One of the most important determinants is the moment of application. Our best results were obtained between 20 to 30% female flower receptivity. Applications at 10% receptivity resulted in less PFA control, probably due to the long bloom period occurring in some areas, and some seasons. This treatment was commercially used in more than 1,000 hectares in Chile, changing 'Serr' production in Chile dramatically. Additionally, others chemicals are in evaluation under Chilean conditions, in order to manage this disorder.

Allelic interaction at seed-shattering loci in the genetic backgrounds of wild and cultivated rice species

It is known that the common cultivated rice (Oryza sativa) was domesticated from Asian wild rice, O. rufipogon. Among the morphological differences between them, loss of seed shattering is one of the striking characters specific for the cultivated forms. In order to understand the genetic control on shattering habit, QTL analysis was carried out using BC(2)F(1) backcross population between O. sativa cv. Nipponbare (a recurrent parent) and O. rufipogon acc. W630 (a donor parent). As a result, two strong QTLs were detected on chromosomes 1 and 4, and they were found to be identical to the two major seed-shattering loci, qSH1 and sh4, respectively. The allelic interaction at these loci was further examined using two sets of backcross populations having reciprocal genetic backgrounds, cultivated and wild. In the genetic background of cultivated rice, the wild qSH1 allele has stronger effect on seed shattering than that of sh4. In addition, the wild alleles at both qSH1 and sh4 loci showed semi-dominant effects. On the other hand, in the genetic background of wild rice, non-shattering effects of Nipponbare alleles at both loci were examined to inspect rice domestication from a viewpoint of seed shattering. It was serendipitous that the backcross plants individually having Nipponbare homozygous alleles at either shattering locus (qSH1 or sh4) shed all the seeds. This fact strongly indicates that the non-shattering behavior was not obtained by a single mutation in the genetic background of wild rice. Probably, some other minor genes are still associated with the formation or activation of abscission layer, which enhance the seed shattering.

異なる標高に生育するコナラとクヌギの離層発達と落葉に関する研究

We investigated the relation between the formation of abscission layer and the leaf shedding of Quercus serrata Thunb. ex. Muuray (the height of test plants were more than 4m, which standing at the height of 80m, 260m, 390m, 630m, 820m above sea level) and Quercus acutissima Carrutherus (the height of test plants were more than 6m, which standing at the height of 120m, 250m, 620m above sea level) on the different altitudes at the south side slope of Mt. Izumi-Katsuragi at Wakayama Prefecture, Japan. The measurement items were the chlorophyll content by SPAD value, the percentage of leaves left at the sampled shoot and photomicrographs of the sliced base of petioles. We measured them every two weeks from October 14, 2005 to January 24, 2006. As a result, the periods of autumn leaves, the formation of abscission layer and the leaf shedding on both species were advanced as the standing levels rising. But some leaves were not shed whether the abscission layer enough developed or not. And, those leaves were forcibly shed at the petiole by the strong wind. The factor of this phenomenon was decrease in strength of the petiole because of drying.

An SNP Caused Loss of Seed Shattering During Rice Domestication

Loss of seed shattering was a key event in the domestication of major cereals. We revealed that the qSH1 gene, a major quantitative trait locus of seed shattering in rice, encodes a BEL1-type homeobox gene and demonstrated that a single-nucleotide polymorphism (SNP) in the 5' regulatory region of the qSH1 gene caused loss of seed shattering owing to the absence of abscission layer formation. Haplotype analysis and association analysis in various rice collections revealed that the SNP was highly associated with shattering among japonica subspecies of rice, implying that it was a target of artificial selection during rice domestication.

ANATOMICAL STUDIES OF THE ABSCISSION PROCESS IN THE JUNCTURE TISSUE BETWEEN THE PEDUNCLE AND SPUR ON JAPANESE PEAR FRUIT FOLLOWING ETHEPHON APPLICATION

ISHS International Symposium on Asian Pears, Commemorating the 100th Anniversary of Nijisseiki Pear ANATOMICAL STUDIES OF THE ABSCISSION PROCESS IN THE JUNCTURE TISSUE BETWEEN THE PEDUNCLE AND SPUR ON JAPANESE PEAR FRUIT FOLLOWING ETHEPHON APPLICATION

Association between Grain Shattering Habit and Formation of Abscission Layer Controlled by Grain Shattering gene sh-2 in Rice(Oryza sativa L.).

Rice garin shattering habit is closely associated with the abscission layer formed at the base of sterile lemmas (empty glumes). In the current study, the character expression of sh-2, one of the representative shattering genes, was investigated by comparing some agronomic parameters among Norin 29, a Japanese non-shattering cultivar, and its shattering near-isogenic line SH-AJNT, as well as Dee-geo-woo-gen, a shattering indica cultivar. After observation of longitudinal sections of sterile lemmas bases, the abscission layer was not found at the spikelet differentiation stage. However, at the reproductive cell formation stage, SH-AJNT faintly formed the abscission layer. No abscission layer, on the other hand, appeared in Norin 29 throughout the stage of panicle development. The breaking strength at the base of sterile lemmas was measured to compare the threshability among materials. Until 19 days after heading, both threshability on Norin 29 and SH-AJNT exhibited similarly high values, indicating non-shattering conditions. At 21 days after heading, the garin of SH-AJNT dropped more easily than those of previou stages, while Norin 29 kept its persistent character. This difference is important for the evaluation of SH-AJNT and Norin 29, shattering and non-shattering cultivars, respectively. The progressive decline in the breaking strength of SH-AJNT was thought to be associated with the ripening of the grain.