1. Introduction1.导言
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Most agricultural crops require pollination (through insects or wind) for successful production. Weather events or asynchrony in flowering between a crop and its polliniser can cause pollination failure, and insect-pollinated crops are additionally vulnerable to declines in insect pollination services. The majority of insect pollination to crops is provided by managed Western honey bees (Apis mellifera) and, for some crops, the regional demand for pollination services can be extreme during bloom, requiring bees to be transported over large distances to service the crop. Globally, pollinator-dependent crops represent an increasing proportion of total agricultural area [1,2], increasing demand on the supply of honey bees and pollination services. While the number of honey bee colonies has increased across the globe in response, the growth rate has not kept pace with demand [3], leading to pollination deficits [4] and increased prices for pollination services. In addition, a number of intersecting issues threaten hive supply during the pollination window, including: increasingly frequent and severe weather events (such as fires, floods, and cyclones) destroying hives as well as resources in the landscape necessary to sustain them [5]; restrictions limiting transport of hives between regions [6,7]; reduction in colony resilience due to pesticide exposure [8]; and competition for hives among crop producers [9]. Sustainable production of high-value crops may require new approaches to pollination services that supplement or substitute honey bees if, or when, there are not enough colonies available to meet demand.
▲猕猴桃针管授粉
大多数农作物需要授粉（通过昆虫或风）才能成功生产。天气事件或作物与其传粉者开花不同步会导致授粉失败，昆虫授粉的作物也容易受到昆虫授粉服务下降的影响。作物的大部分昆虫授粉是由有管理的西方蜜蜂（Apis mellifera）提供的，对于某些作物，在开花期间，该地区对授粉服务的需求可能非常大，需要将蜜蜂长途运输来为作物服务。在全球范围内，依赖传粉媒介的作物在农业总面积中所占的比例越来越大[1,2]，对蜜蜂供应和授粉服务的需求也在增加。尽管全球蜜蜂群体的数量有所增加，但增长速度并没有跟上需求的步伐，导致授粉不足和授粉服务价格上涨。此外，在授粉窗口期间，许多交叉问题威胁着蜂箱的供应，包括：日益频繁和严重的天气事件（如火灾、洪水和飓风）破坏了蜂箱以及维持蜂箱所需的景观资源；限制蜂箱在地区之间运输的限制；农药暴露导致菌落弹性降低；以及作物生产者之间对蜂箱的竞争。如果没有足够的蜂群来满足需求，高价值作物的可持续生产可能需要新的授粉服务方法来补充或替代蜜蜂。
▲Bee pollination
Humans have tried to replace natural pollinators before, with varying degrees of success. Date palms have successfully been hand-pollinated by humans for over 4000 years—documented since at least the Code of Hammurabi [10]. Other plants have been more challenging, as most have smaller flowers, produce less pollen, and have a shorter working life for each pollen grain. Pollen from many plant species has been successfully collected and stored for breeding new varieties, but this requires a tiny fraction of the amount of pollen required to fully replace, or even supplement, insect pollinators [11]. Beyond date palm, the economics of only a handful of other crops, most notably vanilla, can support the labour-intensive cost of pollination by hand. Even in cases where manual pollination has been successful—such as the human pollinators of Maoxian County in China, where a massive force of labourers hand-pollinated vast tracts of apples with paintbrushes in 2001 [12]—they are not always sustainable. Researchers visiting Maoxian Country 10 years later found most of the apple trees had been cut down and replaced by wind-pollinated walnut and self-fertile loquat; both can be pollinated with few, or no, insects [12]. Manual pollination in Maoxian County arose because the high use of pesticides created an environment where beekeepers refused to risk their hives. It was abandoned due to labour migration into the cities.
▲Kiwifruit male flowers
人类以前曾试图取代自然传粉者，取得了不同程度的成功。椰枣树已经成功地由人类人工授粉了4000多年，这至少可以追溯到《汉谟拉比法典》。其他植物则更具挑战性，因为大多数植物的花朵较小，产生的花粉较少，每个花粉粒的工作寿命也较短。许多植物物种的花粉已被成功收集和储存，用于培育新品种，但这只需要完全替代甚至补充昆虫传粉者所需花粉量的一小部分。除了椰枣，只有少数其他作物的经济效益，尤其是香草，可以支持人工授粉的劳动密集型成本。即使在人工授粉成功的情况下——比如中国茂县的人类授粉者，2001年，大量工人用画笔为大片苹果人工授粉——它们也并不总是可持续的。10年后，访问茂县的研究人员发现，大多数苹果树已被砍伐，取而代之的是风媒授粉的核桃和自育枇杷；两者都可以用很少或没有昆虫授粉[12]。茂县的人工授粉之所以兴起，是因为大量使用杀虫剂创造了一个养蜂人拒绝冒蜂箱风险的环境。由于劳动力向城市迁移，它被废弃了。
For several decades, a handful of industries—primarily stonefruit, pipfruit, and kiwifruit—successfully collected and applied pollen using a wide array of pollination technologies, from hand applicators through to tractor-mounted spray systems. Kiwifruit is a flagship crop for artificial pollination, where its dioecious nature (male and female flowers on separate plants), lack of nectar, and high potential for fruit set led to early exploration of artificial pollination [13]. This research demonstrated that supplementing insect pollinators with artificial pollination increased both fruit set and fruit quality [14,15]. The success of early trials in New Zealand led to the development of an array of pollen delivery devices that reduced labour and made artificial pollination an important tool for many growers, and it has become the primary strategy used in Italy [16], southern China [17,18], and Korea [19]. Elsewhere, it is used as a supplemental pollination strategy, with approximately half the kiwifruit growers in New Zealand using supplemental artificial pollination. Several New Zealand companies commercially harvest and mill kiwifruit pollen, supplying it to domestic and global growers [20,21]. Tools developed for kiwifruit have been adapted for, and employed in, other cropping systems [22,23,24].
▲刚完成授粉的红心猕猴桃幼果
▲Kiwifruit male flowers
▲Kiwi insect pollination
Most artificial pollination technologies require a high-quality source of pollen to be successful—a chicken-and-egg situation for crop producers in places that lack existing pollen harvesting and supply infrastructure, although a number of small-scale pollen collection methodologies exist. Despite these limitations, the last decade has seen many researchers, as well as small and start-up companies, take on the pollination challenge, evidenced by the increasing numbers of patents published regarding artificial pollination devices (Figure 1 and Figure 2; Appendix A).
▲猕猴桃鲜果采摘装车
大多数人工授粉技术需要高质量的花粉来源才能成功，这对缺乏现有花粉采集和供应基础设施的地方的作物生产者来说是一种鸡和蛋的局面，尽管存在一些小规模的花粉采集方法。尽管存在这些局限性，但在过去的十年里，许多研究人员以及小型和初创公司都面临着授粉挑战，这可以从越来越多的人工授粉设备专利中得到证明（图1和图2；附录A）。
▲Kiwi insect pollination
2.花粉采集
The quantity of pollen required for artificial pollination depends on both the crop species and the device used to deliver it. Some crops, such as date palm and anemophilous (wind-pollinated) trees, produce large quantities of pollen that is relatively easy to collect. However, most crops, including almond, apple, and kiwifruit, produce a relatively small number of pollen grains per flower and require more labour [29]. This suggests two goals for practical artificial pollination: securing a supply of inexpensive, high-quality pollen, and ensuring that as little pollen as possible is wasted during application.
▲Kiwi electric pollination machine using lithium batteries
人工授粉所需的花粉量取决于作物种类和用于输送花粉的设备。一些作物，如椰枣和风媒（风媒授粉）树，会产生大量相对容易收集的花粉。然而，大多数作物，包括杏仁、苹果和猕猴桃，每朵花产生的花粉粒数量相对较少，需要更多的劳动力[29]。这为实用的人工授粉提出了两个目标：确保廉价、高质量的花粉供应，并确保在应用过程中浪费尽可能少的花粉。
▲猕猴桃人工授粉方式
Pollen is typically most viable at the time of anthesis [30], just as the anthers begin to release pollen. However, collecting flowers at this stage can result in significant pollen losses, as flowers can release the majority of their pollen on the day of anthesis [15,28]. To obtain the highest quantity of pollen per flower, it is generally preferable to harvest immediately prior to anthesis to prevent pollen losses [30].
花粉通常在开花时最具活力[30]，就像花药开始释放花粉一样。然而，在这个阶段收集花朵可能会导致花粉大量流失，因为花朵可以在开花当天释放大部分花粉[15,28]。为了获得每朵花的最高花粉量，通常最好在开花前立即收获，以防止花粉损失[30]。
▲贵州高山猕猴桃种植喜获丰收
Three approaches have been taken to collecting pollen: hand-picking or mechanically harvesting flowers just prior to anthesis and extracting the pollen; vacuuming pollen directly from flowers; and using pollen traps attached to honey bee hive entrances to obtain bee-collected pollen from the crop (Figure 3).
收集花粉有三种方法：在开花前手工采摘或机械收割花朵并提取花粉；直接从花中吸出花粉；并使用附着在蜜蜂蜂箱入口的花粉陷阱从作物中获取蜜蜂采集的花粉（图3）。
Hand-picking individual flowers, inflorescences, or panicles has been successful in plants that produce copious quantities of pollen. For date palm, entire panicles are harvested, dried, and then filtered [28]. The tassels, strobili, or catkins (male reproductive structures) of wind-pollinated plants can be handled similarly [24,31,32,33,34,35].
在产生大量花粉的植物中，手工采摘单个花朵、花序或圆锥花序是成功的。对于椰枣，整个穗被收获、干燥，然后过滤[28]。风媒传粉植物的流苏、球果或柳絮（雄性生殖结构）可以类似地处理[24,31,32,33,34,35]。
▲猕猴桃人工授粉方式
Entomophilous plants (pollinated by insects) typically produce less pollen, making collection more challenging. Anthers can be manually excised from individual flowers—an approach employed in artificial pollination research [36,37], but the method is labour-intensive and is economical only for small-scale application (e.g., breeding programmes, vanilla and cacao pollination). Larger-scale pollination was enabled by methods of mechanically separating pollen from whole flowers for kiwifruit [15,23,38,39], cherimoya [30], date palm [22,28], and almond [40]. This typically involves drying the collected flowers, milling the flowers to separate the anthers from other floral structures, allowing the anthers to dehisce under controlled conditions, and using a filter- or cyclone-type cleaning machine that extracts the anthers and pollen, which can then be dried. The pollen is then extracted using a sieve attached to a vacuum. Anther-drying has been identified as a rate-limiting step in the pollen-milling process outlined above, but attempts to reduce drying time by increasing temperature or air-flow negatively affect viability beyond a certain point [15].
▲Kiwifruit male flowers
昆虫学植物（由昆虫授粉）通常产生的花粉较少，这使得采集更具挑战性。花药可以从单朵花上人工切除——这是人工授粉研究中采用的一种方法[36,37]，但这种方法劳动密集，仅适用于小规模应用（如育种计划、香草和可可授粉）。通过机械分离猕猴桃[15,23,38,39]、番荔枝[30]、椰枣[22,28]和杏仁[40]全花花粉的方法，实现了更大规模的授粉。这通常涉及干燥收集的花朵，研磨花朵以将花药与其他花结构分离，使花药在受控条件下开裂，并使用过滤器或旋风式清洁机提取花药和花粉，然后可以干燥。然后用真空筛提取花粉。花药干燥已被确定为上述花粉碾磨过程中的限速步骤，但试图通过提高温度或空气流量来减少干燥时间，会对超过某一点的存活率产生负面影响[15]。
▲猕猴桃人工授粉方式
Flowers are commonly sourced from growers (either to apply back to the same orchard, or sold for pollen processing), and increasingly, from orchards grown specifically to supply pollen. There are, however, several challenges with orchards dedicated to pollen production, including lower financial return than fruiting orchards and a very brief, high-labour demand for harvesting flowers. As a result, a combination of dedicated pollen production and fruiting orchards is likely to remain in use [41,42,43]. Anecdotally, harvest rates for manual picking vary wildly with the experience of pickers and orchard management practices. A mixed-sex kiwifruit orchard (not dedicated to pollen production), for example, could yield between 20 and 200 kg-flower/ha, with a typical yield of 30–40 kg-flower/ha from a well-trained team in a well-managed orchard. Picking rates vary from less than 1 kg-flower/day up to 40 kg-flower/day, although again this is heavily dependent on experience and orchard practice. Pollen yield for good male cultivars reportedly varies from 8.5 g/kg-flower up to 10.5 g/kg-flower with careful attention to the process. Poor-producing cultivars may yield only 4 g/kg-flower (pers. comm. Mat Johnston).
花卉通常来自种植者（要么用于回用同一果园，要么用于花粉加工），而且越来越多地来自专门为供应花粉而种植的果园。然而，专门用于花粉生产的果园存在一些挑战，包括比果园更低的经济回报，以及收获花朵的劳动力需求非常短暂和高。因此，专门的花粉生产和果园很可能会继续使用[41,42,43]。据传，人工采摘的收获率因采摘者的经验和果园管理实践而异。例如，一个混合性别的猕猴桃园（不专门用于花粉生产）每公顷可以生产20到200公斤的花，在管理良好的果园里，训练有素的团队每公顷的典型产量为30到40公斤的花。采摘率从每天不到1公斤花到每天40公斤花不等，尽管这在很大程度上取决于经验和果园实践。据报道，良好雄性品种的花粉产量从8.5克/千克花到10.5克/千克花不等，要仔细注意这一过程。产量低的品种可能只生产4克/公斤的花（通讯员马特·约翰斯顿）。
▲猕猴桃人工授粉方式
Recently, a mechanical pollen harvesting device has been developed for almond, which uses a tree shaker to dislodge flowers at full bloom, collecting them on a tarp beneath the tree [40,42]. Flowers are dislodged at various states of maturity, so pollen recovery rates are lower on a per-flower basis, but bulk harvesting significantly reduces the cost of collection. Harvesting machinery has also been developed for maize flowers, removing tassels for later milling and application in the production of hybrid maize seed [35]. More than 80 L of pollen can be collected each day with this machinery, a high yield made possible by the fact that maize is wind-pollinated and therefore produces large quantities of pollen [35].
最近，一种用于杏仁的机械花粉采集装置已经开发出来，该装置使用树木摇床在盛开时移走花朵，将它们收集在树下的防水布上[40,42]。花朵在各种成熟状态下都会脱落，因此每朵花的花粉回收率较低，但批量收获显著降低了采集成本。还开发了玉米花收割机械，去除穗以备后续碾磨，并应用于杂交玉米种子的生产[35]。使用这种机器每天可以收集超过80升的花粉，玉米是风媒传粉的，因此可以产生大量花粉，从而实现高产[35]。
Mechanical harvesting techniques are relatively new, so the primary driver of pollen cost is labour. In November 2022, during the pollination season, the price was NZD 8250/kg (USD 5200/kg) for kiwifruit pollen in New Zealand, and USD 7500–8500/kg for pipfruit and stonefruit pollen from the USA [44]. In contrast, date pollen is available at USD 150–225/kg in the USA and Mexico [28].
▲猕猴桃人工授粉方式
机械收割技术相对较新，因此花粉成本的主要驱动力是劳动力。2022年11月，在授粉季节，新西兰的猕猴桃花粉价格为8250新西兰元/公斤（5200美元/公斤），美国的梨果和石果花粉价格为7500-8500美元/公斤[44]。相比之下，在美国和墨西哥，枣花粉的价格为150-225美元/公斤[28]。
In the pursuit of less-expensive material for artificial pollination, there has been considerable interest in pollen collected by honey bees, despite its variable viability and purity. The impurities stem from two sources: honey bees mix pollen with nectar to form pollen pellets on their legs, and additionally may not be foraging exclusively on the target crop. Trials have successfully demonstrated fruit set using bee-collected pollen in kiwifruit [45,46,47], peach [48], apple [48,49], pear [48,50], almond [51,52], and canola [53]. Researchers found, however, that fruit drop was higher and fruit weight lower in flowers fertilised with bee-collected pollen instead of pure pollen. Consequently, higher application rates are required. Several studies have noted lower fruit set from bee-collected pollen, which was improved by washing it and formulating it to remove excess sugars, which otherwise appear to inhibit pollen germination [30,49,53]. A recent study found that pollen on bees’ bodies is much more able to achieve fruit set than that stored on their corbiculae; bees that do not pack their pollen with sugars (e.g., Megachile rotundata, Megachilidae; Halictus spp., Halictidae) do not show the same detrimental effect on pollination potential [53], but it has not yet been possible to collect pollen from these species at sufficient scale. More research into the handling and processing of bee-collected pollen is needed before it can be used for large-scale pollination.
为了寻找更便宜的人工授粉材料，人们对蜜蜂收集的花粉产生了相当大的兴趣，尽管其存活率和纯度各不相同。这些杂质来自两个来源：蜜蜂将花粉与花蜜混合，在腿上形成花粉粒，而且可能不会只在目标作物上觅食。试验已成功证明使用蜜蜂采集的花粉在猕猴桃[45,46,47]、桃[48]、苹果[48,49]、梨[48,50]、杏仁[51,52]和油菜[53]中坐果。然而，研究人员发现，用蜜蜂采集的花粉而不是纯花粉受精的花朵，落果率更高，果实重量更低。因此，需要更高的申请率。几项研究发现，蜜蜂采集的花粉坐果率较低，通过清洗和配制花粉以去除多余的糖来改善坐果率，否则这些糖似乎会抑制花粉萌发[30,49,53]。最近的一项研究发现，蜜蜂身上的花粉比储存在球茎上的花粉更容易坐果；不将花粉包裹在糖中的蜜蜂（例如，圆斑大蜂、圆斑大蚊科、Halictus spp.、Halictidae）对授粉潜力没有同样的不利影响[53]，但还不可能以足够的规模从这些物种中收集花粉。在将蜜蜂采集的花粉用于大规模授粉之前，需要对其处理和加工进行更多的研究。
▲刚完成授粉的红阳猕猴桃幼果
Pollen can also be vacuumed directly from some flowers, catkins, or strobili. The viability of pollen collected in this way is generally much higher than that with other methodologies [30,38]. Furthermore, in some designs, the vacuum may be reversed to apply collected pollen immediately without further processing or storage. Vacuum collection has successfully been employed to collect quantities of larch [54], Douglas fir [55], olive (manual [23], mechanised [30]), kiwifruit [23,38,56], and cannabis [57] pollen. Pollen yield varies with plant species, with yields 500 cc/h being reported for Douglas fir [55], 140 g/h being reported in kiwifruit [38] and 100–200 g/h in olive (cultivar-dependent [23]). A handful of prototype mechanical pollen-harvesting systems have been developed to vacuum pollen from trees. These rely on surrounding the tree with a vacuum and filtration system, while using a tree shaker to release the pollen (olive [30], larch [54]). These mechanised systems can harvest up to ten times the quantity of pollen as handheld vacuum devices can, although the pollen tends to be of somewhat lower quality [54].
花粉也可以直接从一些花、柳絮或球果中抽出。以这种方式收集的花粉的存活率通常远高于其他方法[30,38]。此外，在某些设计中，可以颠倒真空状态，立即施加收集到的花粉，而无需进一步加工或储存。真空收集已成功用于收集大量落叶松[54]、花旗松[55]、橄榄（手动[23]、机械化[30]）、猕猴桃[23,38,56]和大麻[57]花粉。花粉产量因植物种类而异，道格拉斯冷杉的产量为500毫升/小时[55]，猕猴桃的产量为140克/小时[38]，橄榄的产量为100-200克/小时（取决于品种[23]）。已经开发了一些原型机械花粉采集系统，用于从树木中真空采集花粉。这些依赖于用真空和过滤系统包围树木，同时使用树木摇床释放花粉（橄榄[30]，落叶松[54]）。这些机械化系统可以收获的花粉量是手持真空设备的十倍，尽管花粉的质量往往较低[54]。
Different processes are explored in Figure 3.
After collection and processing, binucleate pollen can be put into cold storage and kept viable, often for several years. Storage conditions for particular species have been explored thoroughly in the literature on germplasm maintenance and plant breeding (recently reviewed by [58]). As long as the cold chain is maintained, it is not uncommon to be able to store pollen at −20 °C for 1–6 years with little loss of viability [23,28,58], giving producers flexibility to apply pollen from one year to the next, helping to mitigate synchronicity and production risks.
图3探讨了不同的过程。
经过收集和处理后，双核花粉可以冷藏保存，通常可以保存数年。在种质资源维护和植物育种的文献中，对特定物种的储存条件进行了深入探讨（最近由[58]综述）。只要冷链得以维持，能够在-20°C下储存花粉1-6年而几乎不损失活力的情况并不罕见[23,28,58]，这使生产者能够灵活地将花粉从一年应用到下一年，有助于降低同步性和生产风险。
3. Pollen Application3.花粉施用
Many methods to pollinate plants have been explored. Broadly, there are two main approaches: pollen can be applied dry (possibly diluted with a neutral carrier, such as charcoal, to help manage application rates), or wet (generally suspended in an aqueous liquid, often with additives to ensure isotonic balance with pollen cells) (Figure 4). A third option is available for selected, self-compatible crops—such as tomato—where vibrating the floral structures, either through direct contact or with a puff of air, shakes loose pollen to fertilize the flower [59].
人们已经探索了许多给植物授粉的方法。从广义上讲，有两种主要方法：花粉可以干施（可能用中性载体如木炭稀释，以帮助控制施用量），也可以湿施（通常悬浮在水性液体中，通常含有添加剂以确保与花粉细胞的等渗平衡）（图4）。第三种选择适用于选定的自兼容作物，如番茄，通过直接接触或一股空气振动花朵结构，摇动松散的花粉使花朵受精[59]。
Most application literature measures results directly in terms of fruit or seed set, as pollen requirements have been characterized for relatively few crops [60,61,62,63]. Research has shown that liquid-carrier (or dry diluent composition) formulation, temperature, time of day, weather, stigma receptivity, and flower age all influence pollination efficacy [41,61,64,65,66,67,68]. These factors are largely ignored in much of the work published on alternative pollination systems.
大多数应用文献的测量结果直接与果实或结实有关，因为花粉需求的特征是相对较少的作物[60,61,62,63]。研究表明，液体载体（或干稀释剂组合物）配方、温度、时间、天气、柱头接受性和花龄都会影响授粉效果[41,61,64,65,66,67,68]。在许多关于替代授粉系统的研究中，这些因素在很大程度上被忽视了。
Dry application has two important advantages over wet pollen application:
干施比湿施花粉有两个重要优点：
Dry pollen delivered to non-target areas (e.g., petals, leaves) often remains viable and can be redistributed to the stigmas by bees [69], while pollen remains viable only for a short time once it becomes wet (30–100 min; [15,70]);
递送到非目标区域（如花瓣、叶子）的干花粉通常仍然存活，并可以被蜜蜂重新分配到柱头[69]，而花粉一旦变湿，只能在短时间内存活（30-100分钟；[15,70]）；
Suitable aqueous carrier solutions have been demonstrated for kiwifruit [36], cherry [70], pear [71,72], pistachio [32], and date palm [28], but new solutions must be created, or customized, and validated for each crop.
猕猴桃[36]、樱桃[70]、梨[71,72]、开心果[32]和椰枣[28]已经证明了合适的水性载体溶液，但必须为每种作物创造、定制和验证新的溶液。
Insect pollinators take advantage of naturally occurring electrostatic forces, which help transfer pollen from bee to flower [73]. Research and commercial application have demonstrated that electrostatically charging dry pollen can improve the proportion captured by the flower by 5- to 10-fold [70,74,75,76]. However, droplets in aqueous systems generally have larger mass, making it more difficult to attach a sufficient charge to the pollen to affect its trajectory.
昆虫传粉者利用自然产生的静电力，这有助于将花粉从蜜蜂转移到花朵[73]。研究和商业应用表明，带静电的干花粉可以将花朵捕获的比例提高5到10倍[70,74,75,76]。然而，水性系统中的液滴通常具有更大的质量，这使得更难在花粉上附着足够的电荷来影响其轨迹。
The main advantages of aqueous systems are:
水性系统的主要优点是：
Enabling more targeted delivery, efficacy during damp conditions where insect pollinators may be scarce [15,23,38];
在昆虫传粉者可能稀缺的潮湿条件下实现更有针对性的递送和疗效[15,23,38]；
The additional liquid mass increases momentum for targeted delivery, reducing the risk the pollen is dispersed by wind.
额外的液体质量增加了靶向递送的动量，降低了花粉被风吹散的风险。
The majority of the artificial pollination systems in Figure 4 are used to supplement available bees, but for some crops, these systems perform adequately by themselves, including kiwifruit [16,19,23,77,78,79,80], olive [23], date palm [28], walnut [33,81], tomato [59], and hybrid maize seed [35].
图4中的大多数人工授粉系统用于补充可用的蜜蜂，但对于某些作物，这些系统本身就可以充分发挥作用，包括猕猴桃[16,19,23,77,78,79,80]、橄榄[23]、椰枣[28]、核桃[33,81]、番茄[59]和杂交玉米种子[35]。
In cases where insect pollinators are abundant, bees perform equivalently or better than artificial pollination for kiwifruit [56,82,83] and kiwiberry [84,85], but when conditions are not optimal, particularly in years where local conditions limit pollinator activity, correctly applied supplemental pollination can increase seed number and fruit size [16,19,23,78,79,80]. Pollen is typically applied in two or more passes through the orchard [86], but a single pass at petal fall has been effective in Italian kiwifruit orchards [23].
在昆虫传粉者丰富的情况下，蜜蜂对猕猴桃[56,82,83]和猕猴桃[84,85]的授粉效果与人工授粉相当或更好，但当条件不是最佳时，特别是在当地条件限制传粉者活动的年份，正确应用补充授粉可以增加种子数量和果实大小[16,19,23,78,79,80]。花粉通常在果园中分两次或多次施用[86]，但在意大利猕猴桃果园中，花瓣落下时单次施用是有效的[23]。
▲Kiwi electric pollination machine using lithium batteries
3.1. Hand-Pollination
Manual pollination with a paintbrush, stick, or pole tipped with a feather-brush is labour-intensive [12,23,41,87]. This cost can be sustained by some high-value crops (Table 1) [11], particularly when pollen does not need to be collected (e.g., tomato), the pollen is produced in abundance and collection costs are low (e.g., date), or the market value of the crop is very high (e.g., vanilla). Where mechanisation is available (even if only in the form of a vibratory wand), it is often more effective and economical than manual pollination [23,59,72,86,87].
3.1. 人工授粉
用画笔、棍子或羽毛刷尖端的杆子进行人工授粉是劳动密集型的[12,23,41,87]。一些高价值作物可以承受这种成本（表1）[11]，特别是在不需要收集花粉（如番茄）、花粉大量生产且收集成本低（如枣）或作物的市场价值很高（如香草）的情况下。在机械化可用的情况下（即使只是以振动棒的形式），它通常比人工授粉更有效、更经济[23,59,72,86,87]。
3.2. Handheld Devices
Handheld devices, such as blowers, sprayers, and vibratory wands, are quicker and easier to use for applying pollen than simple hand tools, such as paintbrushes, significantly reducing labour costs. Within these technologies is a constellation of inventive processes, but we found that four general categories of applicators are commonly used in commercial settings: vibratory wands (particularly for indoor tomato production); pneumatic devices tipped with a brush or feather-brush (for orchard crops); handheld blowers (often utilising commercial leaf blowers as a base component); and modified agricultural sprayers (Table 2). Directed broadcast of pollen using a handheld leaf blower has been estimated to deliver only 1% of pollen to stigmatic surfaces [101]; for dry application, some of the remaining 99% may then be redistributed by bees [69]. Other approaches have been explored, including a bubble gun [102], and a handheld electrostatic pollinator that can both collect and apply pollen [103]. However, both require further research to overcome the technical challenges (e.g., poor targeting and damage to floral structures from coronal discharge) before they could become commercial realities. A different handheld electrostatic pollinator was used successfully for the indoor cultivation of hybrid larch seed, but requires an external pollen supply [31].
3.2. 手持装置
手持式设备，如鼓风机、喷雾器和振动棒，比简单的手动工具（如画笔）更快、更容易用于施加花粉，大大降低了劳动力成本。在这些技术中，有一系列创造性的工艺，但我们发现，在商业环境中通常使用四类施用器：振动棒（特别是用于室内番茄生产）；带刷子或羽毛刷的气动装置（用于果园作物）；手持式鼓风机（通常使用商用吹叶机作为基础组件）；以及改良的农业喷雾器（表2）。据估计，使用手持式吹叶机定向传播花粉只能将1%的花粉输送到柱头表面[101]；对于干施，剩下的99%中的一部分可能会被蜜蜂重新分配[69]。已经探索了其他方法，包括气泡枪[102]和可以收集和施用花粉的手持式静电授粉器[103]。然而，在它们成为商业现实之前，两者都需要进一步的研究来克服技术挑战（例如，靶向性差和冠状放电对花结构的破坏）。另一种手持式静电授粉器已成功用于杂交落叶松种子的室内栽培，但需要外部花粉供应[31]。
3.3. Vehicle-Mounted Devices
Sprayers, blowers, and fans of various kinds have been developed for large-scale pollination (Table 3). These devices require far fewer person-hours than equivalent handheld devices [28,86]. Early trials with apples used spray planes [26], and broadcast tractor sprayers [27]). In kiwifruit, broadcast systems were developed in the 1980s and had significant uptake by growers in the USA, Italy, and New Zealand for supplementing insect pollinators (replacing them in Italy, southern China, and Korea [16,17,18,19]) owing to their lower labour costs than hand application. In the Middle East, several inventions, including directed high-pressure sprayers drawn by tractors [28], have also performed as well as or better than the standard practice of manually pollinating date palm. Most of these devices are made to be towed behind a tractor, but some are also able to be mounted on farm ATVs for increased manoeuvrability, or on autonomous robotic platforms to reduce operator costs (e.g., XAG’s R150, Guangzhou, Guangdong, China, a small, multipurpose autonomous spraying and mowing robot). XAG’s ground-based pollen sprayer traversed a row of an apple orchard in 10 min, while hand pollination would take more than 2 h; however, pollination efficacy was not reported [93]. Broadcast wet sprayers are considered a last resort for pollination, and have lower efficiency in kiwifruit than dry applicators unless high rates of pollen are used [23], and also failed to deposit any pollen on the stigma of the much smaller Japanese plum flowers [14].
3.3. 车载设备
已经开发了各种喷雾器、鼓风机和风扇，用于大规模授粉（表3）。这些设备所需的人时比同等的手持设备少得多[28,86]。早期对苹果的试验使用了喷雾机[26]和广播拖拉机喷雾器[27]）。在猕猴桃中，广播系统是在20世纪80年代开发的，由于其比手工应用更低的劳动力成本，美国、意大利和新西兰的种植者大量采用该系统来补充昆虫传粉者（在意大利、中国南方和韩国取代了它们[16,17,18,19]）。在中东，一些发明，包括拖拉机牵引的定向高压喷雾器[28]，也与人工授粉椰枣的标准做法一样好或更好。这些设备中的大多数都是拖在拖拉机后面的，但也有一些能够安装在农场ATV上以提高机动性，或者安装在自主机器人平台上以降低操作员成本（例如，XAG的R150，中国广东广州，一种小型多用途自主喷洒和割草机器人）。XAG的地面花粉喷雾器在10分钟内穿过一排苹果园，而人工授粉需要2个多小时；然而，没有报道授粉效果[93]。喷洒式湿式喷雾器被认为是授粉的最后手段，除非使用高花粉率，否则在猕猴桃上的效率低于干式喷雾器[23]，而且也没有在小得多的日本梅花的柱头上沉积任何花粉[14]。
▲佳沛在国内销售的进口红心猕猴桃
By their nature, non-targeted systems are wasteful, as pollen grains that settle on leaves, orchard structures, and branches do not fertilise the flower (pollen grains that land on petals, which can be redistributed by bees, are in the minority). To address this, researchers have investigated directed broadcast of electrostatically charged pollen, primarily for almonds, but also for apple, pear, sweet cherry, kiwifruit, and pistachio. Directing the pollen into the canopy reduces the amount of pollen lost to the wider environment, while positively charging the pollen increases attraction to pointed structures (such as styles), increasing pollen deposition [108]. Most of the work investigating electrostatic charge in pollination has involved an Israeli group led by Gan-Mor [74,75,108], and two US groups [109,110]. The application of electrostatic charge has increased pollen deposition on stigmas and improved the fruit set of almond, pistachio, date, apple, cherry, and pear under poor pollination conditions [70,94,110]. In general, for insect-pollinated crops other than kiwifruit and date palm, broadcast pollination is applied in addition to bee pollination, where it may improve fruit or nut set in years where pollination is poor [70,110]. However, there have been few studies on the effects of electrostatically charged pollen application on insect-pollinated crops without the assistance of bees; a trial from the grey literature suggests that the systems are no substitute for insect pollinators—almond yields were 1.3% without pollination, 17% with electrostatic pollination, and over 50% with insect pollination [111].
从本质上讲，非目标系统是浪费的，因为落在叶子、果园结构和树枝上的花粉粒不会给花施肥（落在花瓣上的花粉颗粒是少数，可以被蜜蜂重新分配）。为了解决这个问题，研究人员研究了带静电花粉的定向传播，主要用于杏仁，也用于苹果、梨、甜樱桃、猕猴桃和开心果。将花粉引入树冠可以减少花粉流失到更广泛环境中的量，而带正电的花粉可以增加对尖端结构（如花柱）的吸引力，增加花粉沉积[108]。研究授粉中静电荷的大部分工作都涉及由Gan-Mor领导的以色列小组[74,75108]和两个美国小组[109110]。在授粉条件较差的情况下，施加静电荷增加了柱头上的花粉沉积，并改善了杏仁、开心果、枣、苹果、樱桃和梨的坐果率[70,94110]。一般来说，对于猕猴桃和椰枣以外的昆虫授粉作物，除了蜜蜂授粉外，还应用了广播授粉，这可能会在授粉不良的年份提高果实或坚果的结实率[70110]。然而，在没有蜜蜂帮助的情况下，关于静电花粉施用对昆虫授粉作物的影响的研究很少；灰色文献的一项试验表明，这些系统不能替代昆虫传粉者——不授粉时杏仁产量为1.3%，静电授粉时为17%，昆虫授粉时为50%以上[111]。
Table 3
Representative pollination devices mounted to tractors with and without electrostatic charge capability.
表3
安装在具有和不具有静电荷能力的拖拉机上的代表性授粉装置。
| Mode | General Type | Examples | References |
|---|---|---|---|
| Vibration | Fan | Ventola, Italy | [23,41] |
| Dry | Pollen blowers | QuadDuster, KiwiPollen, New Zealand AirShear, KiwiPollen, New Zealand PollenSmart, PollenSmart, New ZealandATV Applicator, PollenPro, USAScumby, Firman Pollen, USA [112]Palm Tree Pollination Machine, AgroPalms Machinery, Spain | [107][41][113][112][28,114,115] |
| Electrostatic pollen blowers | Home-made devicesEdete | [116][95] | |
| Wet | High-pressure sprayers | Palm Tree Pollination Machine, Agrom Agro Machinery, IL | [28] |
| Fogger/fine mist sprayers | Kiwi Pollen Boom Sprayer, KiwiPollen, New Zealand Spruzz@Polline TR, Gerbaudo, Cuneo, Italy XAG R150, China | [38][23,93] | |
| Electrostatic sprayers | Electrostatic Spraying Systems, Inc., USAOnTarget, On Target Spray Systems, USA Fruit Tower, LectroBlast, USA | [111][117][116][93] |
3.4. Unmanned Aerial Vehicles (UAVs)
Drone-based pollination technology has received considerable attention (Table 4). The use of drones to pollinate crops is an attractive proposition both because drones have a good aesthetic fit for the job—they are airborne pollinators, like bees—and because drone technology has a lower barrier to entry than other forms of robotics [118]. These devices are either directly controlled by a pilot, follow a set path defined by the layout of orchard rows, or utilise a 3-D model of the environment built from an earlier pass by scouting drones [119]. Many drone pollinators are modifications of commercially available drones, particularly those designed for agrichemical sprays (e.g., [51,81,120]), but a number are also being custom-designed for pollination. Several pollination modes are being explored, including aerial broadcast distribution of pollen (Table 4), as well as utilising the drone’s air vortices for pollination directly for hybrid grain production and glasshouse-grown self-compatible crops such as strawberry, tomato, pepper, and eggplant [121,122]. Other approaches have also been prototyped, including a microdrone with a fur pad for direct contact pollination [123], and a drone equipped with a bubble gun [102], but the drawbacks to both approaches—time in the first case and accuracy in the second—may limit their applicability in field situations. Indeed, contact-style drones have been criticised for trying to emulate bees too closely: it is neither practicable nor desirable to create tens of thousands of microdrones to replace a single honey bee colony [124,125], let alone the thousands of colonies used each year for intensive commercial cropping. Aerial broadcast approaches are likely to have the same limitations as ground-based broadcasts, with only a small portion of the pollen dispensed reaching the stigma where it can contribute to pollination.
3.4. 无人机
基于无人机的授粉技术受到了相当大的关注（表4）。使用无人机为作物授粉是一个有吸引力的提议，这既是因为无人机具有很好的美学适合这项工作——它们是空中授粉者，就像蜜蜂一样——也是因为无人机技术的进入门槛比其他形式的机器人低[118]。这些设备要么由飞行员直接控制，要么遵循果园行布局定义的设定路径，要么利用侦察无人机从早期通行中构建的环境三维模型[119]。许多无人机传粉者是对商用无人机的改进，特别是那些专为农用化学品喷雾而设计的无人机（例如[51,8112]），但也有一些是为授粉而定制的。目前正在探索几种授粉方式，包括空中传播花粉（表4），以及利用无人机的空气涡流直接为杂交谷物生产和温室种植的草莓、番茄、辣椒和茄子等自交作物授粉[121122]。其他方法也已经原型化，包括带有毛皮垫的微型飞行器用于直接接触授粉[123]，以及配备气泡枪的无人机[102]，但这两种方法的缺点——第一种方法的时间和第二种方法的准确性——可能会限制它们在现场情况下的适用性。事实上，接触式无人机因试图过于接近蜜蜂而受到批评：制造数万个微芯片来取代一个蜜蜂群落既不可行也不可取[124125]，更不用说每年用于密集商业种植的数千个蜂群了。空中广播方法可能与地面广播具有相同的局限性，只有一小部分分配的花粉到达可以促进授粉的柱头。
Drone pollinators have seen a degree of commercial success in date palm pollination, historically pollinated by hand, by humans scaling the palm trees. As these trees produce copious quantities of pollen, there is little concern about waste from broadcast application [108], and the increase in time savings and improved safety from reducing tree climbing are significant [28,119,120]. Walnut pollination is another emerging area, with positive initial results in multiple countries, producing walnut kernels equivalent to those from wind-pollinated controls [33,81]. Dropcopter is the most well-known organization in this field. They provide pollination services to several crops (Table 4), including apple, cherry, and almond, reporting 53%, 40%, and 94% higher fruit set, respectively, over grower standard methods. However, little independent information about this system’s efficacy (and those of most commercial drone offerings) is available to date.
无人机授粉者在椰枣授粉方面取得了一定程度的商业成功，历史上是由人类攀爬棕榈树手工授粉的。由于这些树木会产生大量的花粉，因此很少有人担心广播应用的浪费[108]，减少爬树可以节省大量时间并提高安全性[28119120]。核桃授粉是另一个新兴领域，在多个国家取得了积极的初步成果，生产的核桃仁与风媒授粉对照的核桃仁相当[33,81]。Dropcopter是该领域最知名的组织。它们为多种作物提供授粉服务（表4），包括苹果、樱桃和杏仁，其坐果率分别比种植者标准方法高53%、40%和94%。然而，迄今为止，关于该系统的功效（以及大多数商用无人机产品的功效）的独立信息很少。
Drones have considerable promise in tall tree crops where their ability to dispense pollen above the canopy is an advantage, with potential applications in conifer breeding [129] as well as wind-pollinated nut crops, which have shown promise in artificial pollination trials, including hazelnut, pistachio, and walnut [25,33,81].
无人机在高大树木作物中具有相当大的前景，它们在树冠上方分配花粉的能力是一个优势，在针叶树育种[129]以及风媒授粉的坚果作物中都有潜在的应用，这些作物在人工授粉试验中显示出希望，包括榛子、开心果和核桃[25,33,81]。
3.5. Robotics and Autonomous Pollination
Autonomous robotic pollinators are often designed to target individual flowers for to minimize waste. Generally, machine vision is used to identify a flower that requires pollination. Broadly, two approaches to delivering pollen have been explored: moving an end-effector close to the flower, or spraying pollen from a distance. Several methods to apply pollen with an end-effector have been explored, including using a robot arm to brush the flower with pollen, touching the flower with a vibrating rod, and delivering a tuned vibration or air-blast to distribute pollen. Aside from brushing pollen, these methods apply only to crops that are self-fertile (e.g., tomato). Pollen sprayed from a distance can be delivered wet or dry.
3.5. 机器人与自主授粉
自主机器人传粉者通常被设计为针对单个花朵，以尽量减少浪费。通常，机器视觉用于识别需要授粉的花朵。从广义上讲，已经探索了两种传递花粉的方法：将末端效应器靠近花朵，或从远处喷洒花粉。已经探索了几种使用末端执行器施加花粉的方法，包括使用机器人手臂用花粉刷花，用振动棒触摸花，以及传递调谐振动或气流来分配花粉。除了刷花粉外，这些方法仅适用于自育作物（如番茄）。从远处喷洒的花粉可以湿送或干送。
Machine-vision systems employing deep learning can locate objects in images with 90%, or higher, accuracy, and with processing rates of up to 100 frames/s [97,130,131,132,133]. Systems employing robotic arms tend to be relatively slow. For example, research on a tomato pollination robot reported pollination speeds of 15–20 s per flower, rates that are not practical for pollination of commercial-scale crops [134,135,136]. Several systems that have already been commercialised are quicker—achieving application rates of about 2–5 s per flower cluster [137]—using several arms to pollinate multiple flowers at once and air-jets to reduce the fine motor control required to position end-effectors directly on a flower. For example, Arugga AI incorporate four sets of nozzles to pollinate high-wire tomato [98]. However, systems employing arms to move an end-effector close to the flower have been used only in greenhouse applications to date (Table 5). Field applications tend to favour spraying systems, delivering a dose of pollen from a vehicle moving through the crop. For example, PowerPollen are running commercial trials with a tractor-mounted boom holding 16 parallel autonomous pollinators for maize. The system mechanically funnels maize silks into the autonomous pollinator’s spray region and deliveries a dose of electrostatically charged dry pollen as the tractor traverses the crop rows [35].
采用深度学习的机器视觉系统可以以90%或更高的精度在图像中定位物体，处理速率高达100帧/秒[9713013132133]。采用机械臂的系统往往相对较慢。例如，对番茄授粉机器人的研究报告称，每朵花的授粉速度为15-20s，这一速度对于商业规模作物的授粉是不切实际的[134135136]。几个已经商业化的系统更快——每个花簇的施用率约为2-5s[137]——使用多个臂同时为多朵花授粉，并使用空气喷射器减少将末端执行器直接放置在花上所需的精细运动控制。例如，Arugga AI结合了四组喷嘴来为高架番茄授粉[98]。然而，迄今为止，使用臂将末端执行器移动到花朵附近的系统仅在温室应用中使用（表5）。田间应用往往倾向于喷雾系统，从穿过作物的车辆中输送一定剂量的花粉。例如，PowerPollen正在进行商业试验，其拖拉机上安装的吊杆上装有16个平行的玉米自主传粉者。该系统将玉米丝机械地输送到自主授粉者的喷洒区域，并在拖拉机穿过作物行时输送一剂带静电的干花粉[35]。
Autonomous robotic pollinators equipped with expert-informed targeting systems could identify and target flowers that produce the best fruit, enabling intelligent pollination services—a possibility suggested by Verdant Robotics’ patented apple pollinator and flower thinner (US11308323B2). Unfortunately, very few data are available about the pollination efficacies of the above devices. The prototype ultrasonic strawberry pollinator was able to perform better than hand pollination [99], while the prototype tomato pollinator performed substantially worse [134]. PowerPollen reported success in hybrid maize production, with a 20% boost in seed yield over current practice [35].
配备有专家信息定位系统的自主机器人传粉者可以识别和定位产生最佳果实的花朵，从而实现智能授粉服务——Verdant Robotics的专利苹果传粉者和花朵稀释剂（US11308323B2）提出了这种可能性。不幸的是，关于上述设备的授粉效率，可用的数据很少。原型超声波草莓传粉者能够比手工授粉表现更好[99]，而原型番茄传粉者的表现要差得多[134]。PowerPollen报道了杂交玉米生产的成功，种子产量比目前的做法提高了20%[35]。
Pollen delivered in an aqueous suspension, sprayed from a moving platform, has demonstrated speeds more practical for intensive commercial cropping [77]. In 2019, during field trials of the autonomous research prototype robot, the machine fully pollinated >670 export-quality “Hayward” kiwifruit, without contribution from insect pollinators, from a platform moving at 2.5 km/h. Key metrics (fruit set, seed count, fruit weight) were comparable to those of insect-pollinated control samples [77]. However, the authors cited high pollen usage (3–5 kg/ha), the associated labour cost to collect the pollen, and relatively slow speed as some of the challenges to be overcome before practical commercial application.
从移动平台喷洒的水悬浮液中输送的花粉，已经证明其速度更适合密集的商业种植[77]。2019年，在自主研究原型机器人的田间试验中，该机器从以2.5公里/小时的速度移动的平台上完全授粉了670多个出口质量的“海沃德”猕猴桃，没有昆虫授粉者的贡献。关键指标（坐果、种子数、果实重量）与昆虫授粉的对照样本相当[77]。然而，作者指出，在实际商业应用之前，需要克服的一些挑战包括花粉使用量高（3-5kg/ha）、收集花粉的相关劳动力成本和相对较慢的速度。
4. Conclusions
Artificial pollination is currently used to supplement insect pollinators for a variety of cropping systems. The rare cases where artificial pollination is used instead of insect pollinators are generally characterized by particular features that make natural pollination difficult, such as a lack of natural pollinators (vanilla, cherimoya, date palm) or dioecy combined with abundant pollen (e.g., kiwifruit, pistachio), or, conversely, are those that are easily self-pollinated by agitation alone (e.g., tomato, strawberry). A variety of devices are being developed and are used commercially to pollinate crops, with increasing focus on drone- and robotics-based solutions.
4.结论
人工授粉目前用于补充各种种植系统的昆虫传粉者。使用人工授粉而不是昆虫授粉的罕见情况通常具有使自然授粉困难的特定特征，例如缺乏天然授粉者（香草、番荔枝、椰枣）或结合大量花粉的雌雄异株（如猕猴桃、开心果），或者相反，是那些仅通过搅拌就容易自花授粉的情况（如番茄、草莓）。正在开发各种设备，并将其用于商业作物授粉，越来越多地关注基于无人机和机器人的解决方案。
However, many challenges remain. Insects are full-service pollinators collecting, transporting and delivering a portion of the pollen they collect while foraging. They reproduce quickly, easily producing thousands of “workers” each year, and thus provide the scale needed for intensive commercial cropping systems. Yet, like many ecosystem services, their contribution to food production is threatened. Most artificial pollination research to date treats pollen as a system input and its delivery as the final output. Pollen is available at commercial scale for a limited number of crops globally, and its availability is driven largely by quantified benefits for supplementing natural pollinators. Little research has been found on the collection, processing and management of pollen for artificial pollination in many crops. Perhaps because of the lack of clear commercial benefit and practical methods for application, harvesting pollen is of limited value for all but the highest-value crops. Nevertheless, the increased activity in research and commercial applications over the last decade signals growing appreciation for the vital role pollination services play in agricultural food production systems.
然而，许多挑战仍然存在。昆虫是全方位服务的传粉者，在觅食时收集、运输和递送部分花粉。它们繁殖迅速，每年很容易产生数千名“工人”，从而为密集的商业种植系统提供了所需的规模。然而，与许多生态系统服务一样，它们对粮食生产的贡献受到了威胁。迄今为止，大多数人工授粉研究都将花粉视为系统输入，将其传递视为最终输出。全球范围内，花粉在商业规模上可用于有限数量的作物，其可用性在很大程度上是由补充天然传粉者的量化益处驱动的。在许多作物中，关于人工授粉花粉的收集、加工和管理的研究很少。也许是因为缺乏明确的商业效益和实用的应用方法，除了价值最高的作物外，采集花粉对所有作物的价值都是有限的。然而，过去十年研究和商业应用活动的增加表明，人们越来越认识到授粉服务在农业粮食生产系统中发挥的重要作用。
▲盛开的猕猴桃花
Conceptualization, M.A.B., M.C. and P.M.: methodology, M.A.B. and P.M.; formal analysis, M.A.B.; investigation, M.A.B. and P.M.; data curation, M.A.B.; writing—original draft preparation, M.A.B.; writing—review and editing, M.A.B., M.C. and P.M.; visualization, M.A.B.; supervision, M.A.B.; project administration, M.A.B.; funding acquisition, M.A.B. All authors have read and agreed to the published version of the manuscript.
概念化，M.A.B.、M.C.和P.M.：方法论，M.A.B和P.M。；形式分析，M.A.B。；调查，M.A.B.和P.M。；数据管理，M.A.B。；写作——初稿编写，M.A.B。；写作——审查和编辑，文学硕士、文学硕士和下午。；可视化，M.A.B。；监督，M.A.B。；项目管理硕士。；所有作者均已阅读并同意手稿的出版版本。
We thank Claudia Adams for running the initial patent search, and Warrick Nelson, Monica Holland, Max Buxton, David Pattemore, Nico Bordes, and Anne Gunson for providing a critical review of the manuscript. We also thank three anonymous reviewers for their helpful comments on the manuscript.
我们感谢Claudia Adams进行了初步的专利检索，并感谢Warrick Nelson、Monica Holland、Max Buxton、David Pattemore、Nico Bordes和Anne Gunson对手稿进行了批判性审查。我们也感谢三位匿名审稿人对手稿的宝贵意见。
The authors declare no conflict of interest. All authors have participated in the development of artificial pollination technology. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.
作者声明没有利益冲突。所有作者都参与了人工授粉技术的发展。资助者在研究设计中没有任何作用；收集、分析或解释数据；在撰写手稿时；或者在决定公布结果时。
Footnotes
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▲红心猕猴桃种植
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