Examining the spatial distribution of flower thrips in southern highbush blueberries by utilizing geostatistical methods

Flower thrips (Frankliniella spp.) are one of the key pests of southern highbush blueberries (Vaccinium corymbosum L. x V. darrowii Camp), a high-value crop in Florida. Thrips' feeding and oviposition injury to flowers can result in fruit scarring that renders the fruit unmarketable. Flower thrips often form areas of high population, termed "hot spots", in blueberry plantings. The objective of this study was to model thrips spatial distribution patterns with geostatistical techniques. Semivariogram models were used to determine optimum trap spacing and two commonly used interpolation methods, inverse distance weighting (IDW) and ordinary kriging (OK), were compared for their ability to model thrips spatial patterns. The experimental design consisted of a grid of 100 white sticky traps spaced at 15.24-m and 7.61-m intervals in 2008 and 2009, respectively. Thirty additional traps were placed randomly throughout the sampling area to collect information on distances shorter than the grid spacing. The semivariogram analysis indicated that, in most cases, spacing traps at least 28.8 m apart would result in spatially independent samples. Also, the 7.61-m grid spacing captured more of the thrips spatial variability than the 15.24-m grid spacing. IDW and OK produced maps with similar accuracy in both years, which indicates that thrips spatial distribution patterns, including "hot spots," can be modeled using either interpolation method. Future studies can use this information to determine if the formation of "hot spots" can be predicted using flower density, temperature, and other environmental factors. If so, this development would allow growers to spot treat the "hot spots" rather than their entire field.     

Genomic insight into the developmental history of southern highbush blueberry populations

Interspecific hybridization is a common breeding approach for introducing novel traits and genetic diversity to breeding populations. Southern highbush blueberry (SHB) is a blueberry cultivar group that has been intensively bred over the last 60 years. Specifically, it was developed by multiple interspecific crosses between northern highbush blueberry [NHB, Vaccinium corymbosum L. (2n = 4x = 48)] and low-chill Vaccinium species to expand the geographic limits of highbush blueberry production. In this study, we genotyped polyploid blueberries, including 105 SHB, 17 NHB, and 10 rabbiteye blueberry (RE) (Vaccinium virgatum Aiton), from the accessions planted at Poplarville, Mississippi, and accessions distributed in Japan, based on the double-digest restriction site-associated DNA sequencing. The genome-wide SNP data clearly indicated that RE cultivars were genetically distinct from SHB and NHB cultivars, whereas NHB and SHB were genetically indistinguishable. The population structure results appeared to reflect the differences in the allele selection strategies that breeders used for developing germplasm adapted to local climates. The genotype data implied that there are no or very few genomic segments that were commonly introgressed from low-chill Vaccinium species to the SHB genome. Principal component analysis-based outlier detection analysis found a few loci associated with a variable that could partially differentiate NHB and SHB. These SNP loci were detected in Mb-scale haplotype blocks and may be close to the functional genes related to SHB development. Collectively, the data generated in this study suggest a polygenic adaptation of SHB to the southern climate, and may be relevant for future population-scale genome-wide analyses of blueberry.Subject terms: Agricultural genetics, Plant breeding     

Kaolin affects blueberry maggot behavior on fruit

This study assessed the effects of Surround (kaolin) on several behavioral parameters of female blueberry maggot, Rhagoletis mendax Curran (Diptera: Tephritidae). First fruit visited, walking, cleaning, and oviposition behavior were quantified in two-choice and no-choice assays where females encountered Surround-treated and untreated fruit of highbush blueberries (Vaccinium spp.). In two-choice assays, females had a propensity (68%) to first visit untreated blueberries. In two-choice and no-choice tests, number of walking bouts and duration of walking bouts were significantly shorter on the Surround-treated than on untreated fruit. Few oviposition attempts on fruit were observed, irrespective of treatments and assays. Chromameter measurements showed significant modification of the blueberry skin color parameters lightness, hue, and saturation between untreated fruit compared with fruit dipped once or twice in a suspension of Surround. Oviposition trials with field-treated fruit showed that blueberries treated with Surround had fewer oviposition scars than the control, and this was more pronounced with weekly applications of Surround. Uneven coating of the fruit by Surround in field applications may have resulted in higher acceptance rates by flies than in blueberries individually dipped and used in laboratory trials.     

中国蓝浆果生产现状

全面报道了蓝浆果(Vaccinium L.spp.)在中国的生产情况,共分为南、北两大区域.北区是指以吉林省为主的东北地区,南区指长江以南广阔的酸性红壤土地区.果用野生种只分布在东北地区.自20世纪80年代起开始引种欧美栽培品种,共达60余个.在北区以吉林农业大学为主,在南区以江苏省·中国科学院植物研究所为主,采用组培和扦插方法繁殖,南、北两区均各有年产百万株以上苗木的能力.在南、北两区的适宜地区,如吉林、江苏和贵州等省,都已选出各自的适栽品种,栽培后的产量和果实质量分别均可达到国外优良水平,显示了蓝浆果在中国发展的巨大前景.     

Susceptibility of low-chill blueberry cultivars to Mediterranean fruit fly, oriental fruit fly, and melon fly (Diptera: Tephritidae)

No-choice tests were conducted to determine whether fruit of southern highbush blueberry, Vaccinium corymbosum L., hybrids are hosts for three invasive tephritid fruit flies in Hawaii. Fruit of various blueberry cultivars was exposed to gravid female flies of Bactrocera dorsalis Hendel (oriental fruit fly), Ceratitis capitata (Wiedemann) (Mediterranean fruit fly), or Bactrocera cucurbitae Coquillet (melon fly) in screen cages outdoors for 6 h and then held on sand in the laboratory for 2 wk for pupal development and adult emergence. Each of the 15 blueberry cultivars tested were infested by oriental fruit fly and Mediterranean fruit fly, confirming that these fruit flies will oviposit on blueberry fruit and that blueberry is a suitable host for fly development. However, there was significant cultivar variation in susceptibility to fruit fly infestation. For oriental fruit fly, 'Sapphire' fruit produced an average of 1.42 puparia per g, twice as high as that of the next most susceptible cultivar 'Emerald' (0.70 puparia per g). 'Legacy', 'Biloxi', and 'Spring High' were least susceptible to infestation, producing only 0.20-0.25 oriental fruit fly puparia per g of fruit. For Mediterranean fruit fly, 'Blue Crisp' produced 0.50 puparia per g of fruit, whereas 'Sharpblue' produced only 0.03 puparia per g of fruit. Blueberry was a marginal host for melon fly. This information will aid in development of pest management recommendations for blueberry cultivars as planting of low-chill cultivars expands to areas with subtropical and tropical fruit flies. Planting of fruit fly resistant cultivars may result in lower infestation levels and less crop loss.     

Horizontal and vertical distribution of flower thrips in southern highbush and rabbiteye blueberry plantings, with notes on a new sampling method for thrips inside blueberry flowers

The dispersal behavior of flower thrips was studied during two field seasons within blueberry (Vaccinium spp.) plantings in Florida and southern Georgia. A "shake and rinse" technique used to extract thrips from inside the blueberry flowers was not significantly different from the conventional dissecting technique, but the time taken to complete the extraction of thrips was significantly shorter. Overall, the highest concentration of thrips was captured inside the canopy of blueberry bushes. Using a grid of traps to monitor the dispersal of thrips during the blueberry flowering season, we analyzed their dispersion with graphical and analytical methods to determine and describe their distribution within blueberry plantings. Thrips began to form "hot-spots" 5-7 d after bloom initiation. A hot-spot is defined as a large number of thrips concentrated in a small area of the field, whereas the rest of the field has a low population. The behavior of the population inside these hot-spots fit a Gaussian tendency and a regression was conducted to describe this tendency. Green's and Standardized Morisita's indices were used to determine thrips level of aggregation. Results showed significantly aggregated populations of thrips in both years. Formation of hot-spots in blueberry plantings seemed to be random. However, the formation of hot-spots was higher in places where more than seven thrips per day were captured on sticky traps, 5 to 7 d after the bloom begins. With these results, producers will be able to monitor thrips populations and locate and manage hot-spots before they become a more serious a problem on blueberry farms.     

Nectar robbery by bees Xylocopa virginica and Apis mellifera contributes to the pollination of rabbiteye blueberry

Honey bees, Apis mellifera L., probe for nectar from robbery slits previously made by male carpenter bees, Xylocopa virginica (L.), at the flowers of rabbiteye blueberry, Vaccinium ashei Reade. This relationship between primary nectar robbers (carpenter bees) and secondary nectar thieves (honey bees) is poorly understood but seemingly unfavorable for V. ashei pollination. We designed two studies to measure the impact of nectar robbers on V. ashei pollination. First, counting the amount of pollen on stigmas (stigmatic pollen loading) showed that nectar robbers delivered fewer blueberry tetrads per stigma after single floral visits than did our benchmark pollinator, the southeastern blueberry bee, Habropoda laboriosa (F.), a recognized effective pollinator of blueberries. Increasing numbers of floral visits by carpenter bee and honey bee robbers yielded larger stigmatic loads. As few as three robbery visits were equivalent to one legitimate visit by a pollen-collecting H. laboriosa female. More than three robbery visits per flower slightly depressed stigmatic pollen loads. In our second study, a survey of 10 commercial blueberry farms demonstrated that corolla slitting by carpenter bees (i.e., robbery) has no appreciable affect on overall V. ashei fruit set. Our observations demonstrate male carpenter bees are benign or even potentially beneficial floral visitors of V ashei. Their robbery of blueberry flowers in the southeast may attract more honey bee pollinators to the crop.     

蓝浆果花冠形态特征及其与果实横径相关性的分析

对兔眼蓝浆果（Vaccinium ashei Reade）、北方高丛蓝浆果（V.corymbosum L．）和南方高丛蓝浆果（V.corymbosum hybrids）3个品种群及南方高丛蓝浆果实生系的花冠长度、花冠宽度、花冠长宽比和花冠口径大小等特征进行了测定和比较,并研究了花冠形态特征参数与果实横径的相关性。研究结果表明,蓝浆果不同品种群间和同一品种群内不同品种间的花冠形态特征参数差异较大;依据花冠形态特征参数,可将供试的兔眼蓝浆果和南方高丛蓝浆果各品种及南方高丛蓝浆果各实生系初步划分为4种类型,将供试的北方高丛蓝浆果各品种划分为3种类型。此外,只有南方高丛蓝浆果各品种及其实生系的果实横径与花冠宽度相关性显著（P〈0．05）,相关系数分别为0．639和0．324。     

Settling behavior of the glassy‐winged sharpshooter,Homalodisca vitripennis,vector of Xylella fastidiosa,on southern highbush blueberry cultivars

The xylem‐limited bacterium Xylella fastidiosa Wells, Raju, Hung, Wiseberg, Mandelico‐Paul & Brenner causes bacterial leaf scorch, a new disease of southern highbush blueberry [Vaccinium corymbosum L. (Ericaceae) interspecific hybrids] in the southeastern USA. The bacterium is transmitted by sharpshooter leafhoppers (Hemiptera: Cicadellidae), with the glassy‐winged sharpshooter (GWSS), Homalodisca vitripennis (Germar), being the most common potential vector in blueberry plantings. Considerable differences are observed in disease intensity among blueberry cultivars in the field, but it remains unknown whether these differences are due to variations in cultivar susceptibility to the bacterium, differences in attractiveness to its vector(s), or both. The settling and feeding behavior of GWSS was evaluated on three southern highbush blueberry cultivars (FL 86‐19, Star, and Emerald) in the greenhouse; in previous field studies, these cultivars were found to be affected by bacterial leaf scorch at high, moderate, and low levels, respectively. Potted plants were caged together with field‐collected GWSS adults for a period of 5 days, with either all three cultivars caged simultaneously (choice test) or individually (no‐choice test). In the choice tests, no differences in cultivar attractiveness were observed during the first 2 days of study. However, from the 3rd day, significantly more adults settled on Emerald (52.7%) than on Star (25.5%) and FL 86‐19 (17.7%). In contrast, in no‐choice tests, the overall host plant acceptance for the three cultivars was similar, and 89.3–92.1% of adults settled on these plants. A significantly greater number of adults settled on the stems (79.3%) vs. the leaves (11.2%). Moreover, they preferred the middle portion of the shoot (56.9%) compared with the upper (7.8%) and the lower (14.7%) portions. In a separate study on FL 86‐19, greater numbers of GWSS nymphs settled on the upper (actively growing) portion of the shoot (82.3%), whereas only 15.8 and 1.5% settled on the middle and lower portions, respectively.     

Bombus impatiens (Hymenoptera: Apidae): an alternative to Apis mellifera (Hymenoptera: Apidae) for lowbush blueberry pollination.

The pollination effectiveness of the commercially reared bumble bee Bombus impatiens Cresson, was compared in field studies to the honey bee, Apis mellifera L., for lowbush blueberry, Vaccinium angustifolium Ait. A preliminary study indicated that B. impatiens had potential as an alternative pollinator. In a 3-yr study, percentage fruit set, percentage harvested berries, berry weight, and seeds per berry were compared in blueberry fields stocked at 7.5 A. mellifera hives per hectare to 5, 7.5, or 10 B. impatiens colonies per hectare. Percentage of harvested berries (yield) was significantly higher in fields stocked with B. impatiens at 10 colonies per hectare. No other parameters measuring pollinator effectiveness were significantly different at 5, 7.5, or 10 colonies per hectare. Flower handling time was significantly faster for B. impatiens and it more frequently collected blueberry pollen. All parameters of pollinator effectiveness were similar for B. impatiens, A. mellifera, and native wild bees in a follow-up study. Overall, B. impatiens was a suitable alternative to A. mellifera.     

Phenology and infestation patterns of plum curculio (Coleoptera: Curculionidae) on four highbush blueberry cultivars

The plum curculio, Conotrachelus nenuphar (Herbst), is a well known pest in apple and peach orchards, but it also is capable of having an economic impact in highbush blueberries. Host phenology and plum curculio oviposition patterns were determined on four highbush blueberry cultivars differing in fruit maturation period. Numbers of oviposition scars were higher on early- ('Weymouth') and mid-season ('Duke' and 'Bluecrop') blueberries than on late-season 'Elliott' in 2001, 2002, and 2003. In 2002, eggs were first present on the three earliest cultivars 21 d before those on 'Elliott', whereas eggs were found on 'Elliott' >40 d after the last sample with eggs for the other three cultivars. The pattern of host phenology and infestation levels suggested that plum curculio oviposition synchronizes well with the availability of suitable fruit for oviposition on early and mid-season cultivars compared with a late-season cultivar of highbush blueberries. The implications of a transition to use of reduced-risk insecticides are discussed in relation to plum curculio management.     

Commercial bumble bee Bombus impatiens (Hymenoptera: Apidae) as a pollinator in lowbush blueberry (Ericale: Ericaceae) fields

Here, we evaluate the potential of the bumble bee Bombus impatiens Cresson, obtained from commercial colonies, as a pollen vector for lowbush blueberry, Vaccinium augustifolium Aiton. We wanted to gain insight into the relationships between pollination by introduced bumble bees, the categories of seeds produced, and the weight and the maturity of the blueberries. The effect of B. impatiens foraging in blueberry stands was measured quantitatively through fruit set and seed set analysis. We created a density gradient of B. impatiens by clustering 72 small hives at the southern extremity of blueberry fields located in Girardville (49 degrees 00' N, 72 degrees 33' W), Quebec, Canada. Observers recorded plant and insect data in 52 plots of 1 by 10 m, distributed at distances ranging from 25 to 1,500 m from the hives. From these data, we evaluated the fruit set rate, the weight and maturity of the berries, and the number of seed structures per berry, including true seeds of large, medium, and small size, pseudo-seeds, and ovules. Positive correlations were found between the density of B. impatiens and fruit set as well as the number of large seeds per berry. Large seeds influenced the weight and maturity of berries. B. impatiens acted as a "near-nest central forager" and increased fruit set and seed production up to 100 and 150 m respectively, from the hives.     

Influence of native flowering plant strips on natural enemies and herbivores in adjacent blueberry fields

Conservation plantings of native wildflowers were established adjacent to highbush blueberry (Vaccinium corymbosum L.) fields to test the hypothesis that provision of resources for natural enemies increases their abundance in adjacent crop fields without increasing the abundance of pest insects. For two growing seasons, natural enemies and herbivorous insects were sampled in fields with flowering borders and in control fields where growers maintained standard mown grass perimeters. Insects were categorized according to their trophic level and their potential pest status, and their abundance was compared between years and between treatments. Syrphid flies (Diptera: Syrphidae) were significantly more abundant in fields with conservation strips, as were plant bugs (Hemiptera: Miridae), thrips (Thysanoptera: Thripidae), and hoppers (Hemiptera: Auchenorrhyncha). Aphids (Hemiptera: Aphididae), thrips, fruit flies (Diptera: Tephritidae), and pirate bugs (Hemiptera: Anthocoridae) decreased significantly in abundance from 2007 to 2008. Beneficial insect abundance in crop fields increased in the latter half of the season in both years and this increase was more pronounced in fields adjacent to conservation plantings. We discuss the implications of these findings for pest management and conservation of biodiversity in farmland.     

Lethal and sublethal activities of imidacloprid contribute to control of adult Japanese beetle in blueberries

Field-based bioassays and residue profile analysis were used to determine the relative importance of lethal and sublethal effects of imidacloprid on adult Japanese beetle, Popillia japonica Newman, in blueberries, Vaccinium corymbosum L. Field-based bioassays assessed adult mortality and knockdown, and fruit and leaf injury from Japanese beetles exposed to 4-h and 7-d field-aged residues of imidacloprid, and the conventional insecticides azinphosmethyl and esfenvalerate. Azinphosmethyl and imidacloprid caused high levels of mortality when beetles were exposed to blueberry shoots with ripe fruit 4 h postapplication, and all compounds protected blueberry fruit and foliage from beetle feeding. Azinphosmethyl and esfenvalerate caused significant Japanese beetle mortality when adults were exposed to blueberry shoots 7 d postapplication, whereas imidacloprid residues caused effects that protected leaves, although not of ripe fruit. When beetles were exposed to shoots with immature green fruit, relatively more leaf feeding and mortality were observed, suggesting that earlier treatment timings may be most effective for systemic neonicotinoids. Japanese beetle mortality was highly correlated with imidacloprid fruit and leaf surface residues, whereas sublethal feeding deterrent effects were observed after the surface residues diminished. The value of the plant-insect-chemistry model for describing the spatial and temporal dimensions of insecticide modes of activity is discussed in terms of optimizing crop protection.     

Can Mass Trapping Reduce Thrips Damage and Is It Economically Viable? Management of the Western Flower Thrips in Strawberry

The western flower thrips Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) is a cosmopolitan, polyphagous insect pest that causes bronzing to fruit of strawberry (Fragaria x ananassa). The main aim of this study was to test whether mass trapping could reduce damage and to predict whether this approach would be economically viable. In semi-protected strawberry crops, mass trapping of F. occidentalis using blue sticky roller traps reduced adult thrips numbers per flower by 61% and fruit bronzing by 55%. The addition of the F. occidentalis aggregation pheromone, neryl (S)-2-methylbutanoate, to the traps doubled the trap catch, reduced adult thrips numbers per flower by 73% and fruit bronzing by 68%. The factors affecting trapping efficiency through the season are discussed. Damage that would result in downgrading of fruit to a cheaper price occurred when bronzing affected about 10% of the red fruit surface. Cost-benefit analysis using this threshold showed that mass trapping of thrips using blue sticky roller traps can be cost-effective in high-value crops. The addition of blue sticky roller traps to an integrated pest management programme maintained thrips numbers below the damage threshold and increased grower returns by a conservative estimate of £2.2k per hectare. Further work is required to develop the F. occidentalis aggregation pheromone for mass trapping and to determine the best timing for trap deployment. Mass trapping of thrips is likely to be cost-effective in other countries and other high-value crops affected by F. occidentalis damage, such as cucumber and cut flowers.     

Native Vaccinium spp. and Gaylussacia spp. infested by Rhagoletis mendax (Diptera: Tephritidae) in the Great Lakes Region: a potential source of inoculum for infestation of cultivated blueberries.

In this study, we addressed the question of whether or not native stands of blueberry (Vaccinium spp.) and/or huckleberry (Gaylussacia spp.) support populations of blueberry maggot, Rhagoletis mendax Curran, in the Great Lakes region. Infestation of commercial blueberries by the blueberry maggot, R. mendax, is a serious problem in many areas where blueberries are grown. In the past 10-20 yr, commercial bighbush blueberry, Vaccinium corymbosum L., production has expanded into places such as southern Ontario and southern Quebec where blueberry maggot had not previously been reported. In the mid-1990s, isolated infestations of commercial highbush blueberry were reported in southern Ontario. Because R. mendax was not considered endemic to that area, it was widely assumed that the pests had come into the fields via movement from exotic localities. Here we present an alternative hypothesis, that the blueberry maggots infesting newly established highbush plantations are derived from native blueberries growing in the vicinity. To test this hypothesis, in 1997-1999, we sampled potential native hosts for R. mendax (Vaccinium spp. and Gaylussacia spp.) from 31 localities in the Great Lakes region, primarily in Michigan and Ontario. R. mendax was reared from fruits of native hosts collected at four sites in Michigan and one site each in Ontario, Indiana, and Ohio. V. corymbosum was the predominant host infested, with infestation of this host observed at five of the seven sites. However, two huckleberry species [Gaylussacia baccata (Wangenheim) K. Koch, and Gaylussacia dumosa (Andersson) Torrey & Gray] had the highest rates of infestation that we observed (25.4 and 17.6%, respectively). These data represent the first published reports of R. mendax infesting native host plants in the Great Lakes region, and support the hypothesis that infestations observed in commercial fields may have originated from infested native host plants.     

Spotted wing drosophila (Drosophila suzukii) utilization and dispersal from the wild host Asian bush honeysuckle (Lonicera spp.)

相似文献   

Variation in highbush blueberry floral volatile profiles as a function of pollination status, cultivar, time of day and flower part: implications for flower visitation by bees

Background and Aims

Studies of the effects of pollination on floral scent and bee visitation remain rare, particularly in agricultural crops. To fill this gap, the hypothesis that bee visitation to flowers decreases after pollination through reduced floral volatile emissions in highbush blueberries, Vaccinium corymbosum, was tested. Other sources of variation in floral emissions and the role of floral volatiles in bee attraction were also examined.

Methods

Pollinator visitation to blueberry flowers was manipulated by bagging all flowers within a bush (pollinator excluded) or leaving them unbagged (open pollinated), and then the effect on floral volatile emissions and future bee visitation were measured. Floral volatiles were also measured from different blueberry cultivars, times of the day and flower parts, and a study was conducted to test the attraction of bees to floral volatiles.

Key Results

Open-pollinated blueberry flowers had 32 % lower volatile emissions than pollinator-excluded flowers. In particular, cinnamyl alcohol, a major component of the floral blend that is emitted exclusively from petals, was emitted in lower quantities from open-pollinated flowers. Although, no differences in cinnamyl alcohol emissions were detected among three blueberry cultivars or at different times of day, some components of the blueberry floral blend were emitted in higher amounts from certain cultivars and at mid-day. Field observations showed that more bees visited bushes with pollinator-excluded flowers. Also, more honey bees were caught in traps baited with a synthetic blueberry floral blend than in unbaited traps.

Conclusions

Greater volatile emissions may help guide bees to unpollinated flowers, and thus increase plant fitness and bee energetic return when foraging in blueberries. Furthermore, the variation in volatile emissions from blueberry flowers depending on pollination status, plant cultivar and time of day suggests an adaptive role of floral signals in increasing pollination of flowers.     

Management of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), in hydroponic strawberry crops: using yellow sticky traps to determine action thresholds

Abstract  The relationship between western flower thrips (WFT) populations in flowers and catches on yellow sticky traps was investigated in a hydroponic strawberry crop in the Sydney area in 1999–2000. The thrips population was initially dominated by males, but by mid-summer it was composed primarily of females. At a point when approximately 60–65% of WFT on traps were female (approximately 20–30 females per trap), the overall density of thrips on traps and in the crop increased rapidly, resulting in severe damage to immature fruit. The sex ratio of 0.60–0.65 females corresponded very closely to a mean of five adult WFT per flower at the start of the trapping week, which was an infestation level previously calculated as the action threshold to prevent damage to young green fruit.     

我国和世蓝浆果的发展前景

蓝浆果是具独特保健作用、生食加工皆宜的适于我国红黄壤地区开发的新兴经济作物。通过对美国一些研究机构和苗圃的考察及参加在智利举行的第七届国际蓝浆果研讨会,对目前世界蓝浆果的发展状况、趋势及一些最新研究成果作了概述,对我国发展蓝浆果的前景、方向等进行了讨论。