云南草蔻花蜜分泌格局与访花动物行为及其对果实和种子产量的影响

依靠动物传粉的植物, 其繁殖成功 (胚珠受精与花粉散布 ) 很大程度上取决于访花动物的种类和拜访行为, 而访花动物的种类和行为又受植物提供给传粉者的回报, 主要是花蜜分泌格局的影响。通过对姜科山姜属植物云南草蔻 (Alpiniablepharocalyx) 花蜜分泌量及糖分含量的测定, 拜访动物种类、传粉昆虫拜访频率及停留时间的调查, 主要盗蜜者花松鼠 (Tamiopsswinhoei) 的盗蜜行为观察, 植物的结实率以及结籽量的分析等, 探讨花蜜分泌与传粉昆虫的关系以及盗蜜对果实及种子生产的影响。研究表明 :云南草蔻的两种花型花蜜的分泌量在一天的单花开花过程中呈现相反的格局, 花柱上举型花下午的花蜜分泌量高于上午, 而花柱下垂型花则上午略高于下午 ;两种花型花蜜的糖分浓度都随着开花时间而下降 ;共有 17种访花动物拜访云南草蔻的花, 其中 8种具有传粉作用 ;传粉昆虫的停留时间随拜访频率的不断增加而缩短 ;被盗蜜植株与未被盗蜜植株的结实率差异不显著而结籽量差异显著。     

多叶斑叶兰花蜜特征和分泌规律及其对昆虫访花的影响

为探讨多叶斑叶兰(Goodyera foliosa)花蜜体积和成分的变异性,及其对传粉昆虫的影响,采用毛细管法、折光仪和高效液相-蒸发光法对一天内不同时间段单花花蜜体积、总糖浓度和可溶性糖成分及含量进行检测,用摄像机对其传粉昆虫访花行为进行观察。结果表明,单花花蜜体积在10:00达到最大[(7.19±2.29)μL, n=10],糖浓度在16:00达到最大[(25.85±1.83)%,n=10]。花蜜中的主要可溶性糖为果糖和蔗糖,果糖在上午10:00含量达到最高(78.310 mg/g);蔗糖在下午16:00含量达到最高(247.600 mg/g)。中华蜜蜂(Apis cerana)访花时间为每天9:00-15:00,访花高峰期在10:00-14:00;橘尾熊蜂(Bombus friseanus)访花时间为每天8:00-16:00,访花高峰期在10:00-12:00和16:00-18:00。因此,这两种传粉昆虫访花频率与多叶斑叶兰单花花蜜体积及糖浓度的变化有一定的相关性,它们更倾向于访问较高花蜜体积及糖浓度的花朵。多叶斑叶兰花蜜成分中蔗糖占优势,能有效吸引各种蜂类、蝶类和蚁类访花,且访问频率较高,中华蜜蜂和橘尾熊蜂能携带其花粉块,是主要的有效传粉昆虫。花形态和访花昆虫的体型大小的匹配,决定了是否成为有效传粉昆虫。     

贠建全:广东荷包岛维管束植物名录

以分布于秦岭的金花忍冬（Lonicera chrysantha Turcz.）、忍冬（L.japonica Thunb.）、葱皮忍冬（L.ferdinandii Franch.）和金银忍冬（L.maackii（Rupr.）Maxim.）为对象,通过定位观察、人工授粉实验、人为设置实验斑块的方法对忍冬属4种植物的开花生物学特性、繁育系统、花色变化现象、传粉过程进行了研究。结果表明,4种植物的单花花期、花部特征存在差异。人工授粉实验显示,4种植物均存在一定的花粉限制,自交不亲和。除葱皮忍冬外,其余3种植物随着花色由白变黄,花粉和花蜜报酬减少、雌雄生殖能力逐渐降低;葱皮忍冬花变色后花蜜量变化不显著,且仍保留较强的雌性生殖能力。变色花的保留被认为是植物的一种生殖策略,通过增大植物的花展示来扩大自身的吸引力,以吸引更多远距传粉者访花。人为控制白、黄花不同数量比的实验结果表明,大多数传粉者偏向访问白花（变色前的花）,且白花提供的报酬量和黄花（变色后的花）数量显著影响传粉者的访花频率,即当花蜜量减少或黄花数量增多时,传粉者访花频率随之降低。因此,我们认为忍冬属4种植物的花色变化可能除了增大植物对远距传粉者的吸引力外,对近距传粉者的访花行为也可能具有一定的影响。当传粉者接近植株时,变色后的花可能暗示其花蜜（花粉）报酬已经发生变化,并驱使昆虫离开并飞向同株或异株植物新开放的报酬丰富的白花,这既有利于提高传粉者的觅食效率,又能降低植物同株异花授粉的几率,对忍冬属植物及传粉者都具有重要意义,是植物长期与授粉昆虫相互适应的反映。     

刺五加传粉生物学研究

本文报道刺五加Eleutherococcus seticosus(Rupr．et Maxim．)Maxim的花朵酬物、访花者类别、访花 者在花序上的访花行为、访花频率及传粉效果。主要结果如下:(1)每朵雄花可提供的报偿是58 000～ 81 000粒花粉,5.5～8.0 µl／d×2～4d花蜜。每朵雌花仅可提供4.0～8.0µl／d× 2～3 d花蜜,不能提 供花粉。每朵两性花可提供19 000～54 000粒花粉和7.0～10.0 µl／d× 2～6 d花蜜。(2)花粉提供者 (雄花和两性花)提供报偿的日期是开花后1～3天,花粉接受者(雌花和两性花在柱头外翻变白时)是 开花后5～7或7～9天,这进一步证实了剌五加雄蕊先熟。另外,花粉提供者提供花蜜的时间在一天中 是8:30至15:30,其高峰是9:00～15:00;花粉接受者提供花蜜的时间在一天中是10:30至16:30,其高 峰是11:00～16:00。这种时间差异可能是刺五加影响传粉者流向(从花粉提供者到花粉接受者)的关键 因素。(3)在刺五加花朵上记录到的访花昆虫有50余种,分别隶属于膜翅目、鳞翅目、鞘翅目、双翅目和 半翅目。在不同天气、不同生境、不同性别的植株记录到的访花频率及高峰时间不同:花粉提供者的 访花高峰早于花粉接受者的;雄株上的访花频率高于雌株的;两性株接受花粉时的访花频率高于提供花 粉时的;而天气越晴朗,刺五加地块(patch)越大,则访花频率越高;每日接受光照越早,访花高峰越早。 (4)从传粉效果看,刺五加种子的形成完全依赖于传粉昆虫的活动。其中,蜂类,如熊蜂、花蜂、切叶蜂、胡蜂和蜜蜂等是最有效的传粉者,其它昆虫如粉蝶、食蚜蝇、寄蝇和鳃角金龟等对传粉也有帮助。     

八角金盘传粉昆虫及其访花行为

八角金盘(Fatsia japonica(Thunb.)Dccne.et Planch.)为冬季开花植物,探讨其访花昆虫种类及访花行为对研究冬季开花植物的传粉生物学规律具有重要意义。对八角金盘访花昆虫种类、访花频率、访花时间、访花行为进行了初步研究。结果发现:八角金盘的传粉昆虫有2目、3科、3种,主要为膜翅目胡蜂科金环胡蜂Vespa mandarinia Smith、双翅目食蚜蝇科斜斑鼓额食蚜蝇Scaeva pyrastri(L.)及丽蝇科大头金蝇Chrysomya(Compsomyia)megacephala(Fabricius);八角金盘提供的酬物主要为花粉与花蜜,食蚜蝇舔食其花粉与花蜜,而金环胡蜂与大头金蝇主要吮吸花蜜;金环胡蜂、大头金蝇及斜斑鼓额食蚜蝇的访花高峰均在10:30—14:30;斜斑鼓额食蚜蝇访花时间较长,5～10min,金环胡蜂访花时间较短,8～30s,大头金蝇访花时间最短只有3～5s。八角金盘具有较低的自花授粉能力,但主要为异花授粉植物。     

毛菍传粉生物学研究

为探讨毛菍(Melastoma sanguineum Sims.)传粉生物学特性,对海南霸王岭国家级自然保护区林缘、林内和林隙3个生境内毛菍的花期、花部形态、访花昆虫等进行观察,同时对其繁育系统相关指标进行了测定。结果表明：毛菍无花蜜分泌;花瓣对访花昆虫有吸引作用;访花昆虫主要有蜂类、食蚜蝇类和丽金龟类;主要传粉昆虫有木蜂、熊蜂,其中木蜂传粉活动最为活跃,为最有效传粉昆虫。毛菍具典型的异型雄蕊,长、短雄蕊在传粉过程中具有明确的功能分化,推测这是毛菍保证传粉成功的策略。毛菍为自交亲和的异交种,需要昆虫传粉;人工自交和异交均具有较高的坐果率;不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象;其繁育系统是兼性自交。昼夜温差较小和相对湿度较高的林内和林隙生境有利于延长群体花期;光照充足、温度相对较高的林缘生境,花朵发育较快,柱头可授性的时间稍早。花粉、传粉者和环境条件是制约毛菍野外传粉效率的主要因素。     

益智传粉生物学的研究

 益智（Alpinia oxyphylla）的花期从2月底至4月下旬;单株花期（花序）约为16～26 d,通常为23～26 d;单花花期一般为1 d。正常情况下,益智和其它山姜属植物一样,具有花柱卷曲性促进异花授粉的机制,表现出两种类型：花柱上举型和花柱下垂型,其花柱卷曲运动的节律与其它已报道的山姜属植物基本一致。但观察发现,当遭遇低温天气时(日最高气温<18℃),单花花期延长为2 d,无论是上举型个体还是下垂型个体,均只表现出一种花型——下垂型：上午开花时花柱弯向上,柱头位于已开裂散发出花粉的花药上方,直到第二天上午6∶30～11∶00间,花柱才陆续地慢慢向下运动,柱头下降至与花药等高或位于花药下方。益智的主要传粉昆虫是蜜蜂（Apidae sp.）、木蜂（Xylocopa sp.）,绝大多数的访花者的访花目的是吸蜜。益智的花蜜分泌量（8.37～15.79 μl）和花蜜含糖量（30.12%～32.83%）较高,花蜜是益智作为传粉者访花的最主要的报酬。实验结果还表明,益智花部中唇瓣对昆虫访花有显著的招引作用;益智的花蜜对蜜蜂的访花频率有显著的影响,对木蜂有一定的招引作用,但并不显著。而花粉（花药）则对昆虫的访花频率影响不大。人工授粉实验结果表明益智存在自交亲和性,无论是上举型或下垂型个体自交和异交均有较高的结实率;人工自交和异交的结实率在上举型植株中存在较大的差异,而在下垂型个体中则差异不明显;去雄套袋、去柱头套袋和完全套袋不授粉等处理均不结实,表明益智不存在无融合生殖现象和自动自花授粉现象。益智的繁育系统是异花授粉交配系统。     

草果两型植株的繁育差异研究

草果(Amomum tsaoko)是具有柱头卷曲机制的姜科植物,该研究通过观察和分析传粉者的访花行为、草果的花蜜分泌和最终的结实率,探索其花蜜分泌模式与传粉行为的相互关系,以及对其两型植株繁殖的影响。结果表明:草果两型植株的花蜜分泌在1 d花期的16:40—19:00期间达到高峰;上举型花比下垂型花能够提供更多的花蜜,下垂型花的花蜜分泌受环境温度、光照、湿度的显著影响,其结实率也显著低于上举型花。中华蜜蜂(Apis cerean cerana)是观察点频率最高的有效传粉者,在1 d花期里访花频率呈现双峰模式,访花行为受到温度和湿度的显著影响,78%或18℃较适宜访花。1 d花期中温度、湿度的变化,以及花蜜的动态分泌模式共同促进这一双峰访花模式的形成。草果花蜜分泌模式和双峰访花模式均与草果的柱头卷曲运动匹配,对吸引昆虫精确传粉、维持柱头卷曲机制有重要意义。草果两型植株之间,环境因素对它们花蜜分泌的影响显著不同,导致传粉者不同的访花行为反应和繁殖结果。适当增加居群中上举型植株数量,可能是提高产量的一个有效方法。     

濒危植物翅果油树种群传粉生物学研究

对山西省南部平陆、乡宁和翼城3个翅果油树(Elaeagnus mollisDiels)种群分别进行了传粉生物学观察.结果表明:(1)翅果油树花期一般为8~12 d,单花花期3~4 d,结实率仅为0.42%;(2)访花者多属于膜翅目、鞘翅目、双翅目、鳞翅目昆虫,蜂类是有效的传粉者;(3)翅果油树依靠芳香气味、花蜜、花粉和适度紧缩的生殖枝吸引访花昆虫;(4)昆虫的访花活动主要集中在11:00~16:00,访花活动易受气象因素影响;(5)自然条件下,翅果油树平均每日每花被访问的频次为(1.93±1.64)次,人工放蜂可显著增加传粉强度;(6)交配系统属居群内的异花传粉,仍属于广义的自花传粉范畴.传粉昆虫少,传粉效率低是翅果油树致濒的重要因素之一.     

腐生植物无叶美冠兰食源性欺骗传粉研究

无叶美冠兰是一种典型的腐生兰科植物,为揭示该物种的自然传粉机制,拓展对兰科植物生殖特性的认识,在广西雅长兰科植物国家级自然保护区对其开展了传粉生态学观测研究。结果表明:无叶美冠兰花朵具备高度自交亲和能力,但不存在自动自花授粉机制,必须依赖外部传粉媒介把花粉送到柱头,实现有效传粉;绿彩带蜂是无叶美冠兰唯一有效传粉昆虫;传粉昆虫与花朵在与传粉功能相关的关键性状在形态上良好拟合;绿彩带蜂的访花活动主要发生在3个阶段:8.6%发生在9:00~11:30,80.2%发生在11:30~14:00,11.2%发生在14:00~15:30;花朵在中午强烈的阳光直射下挥发出香甜的气味。无叶美冠兰花朵主要通过挥发极具诱惑力的香甜气味和唇瓣上黄色的蜜导来诱导绿彩带蜂进入花朵中觅食,传粉昆虫与花朵在与传粉功能相关的关键性状在形态上良好拟合促成有效传粉,绿彩带蜂在整个传粉过程没有获得报酬,是食源性欺骗传粉机制。     

Deceptive pollination of Calanthe by skippers that commonly act as nectar thieves

Flowers have developed different strategies to attract pollinators through visual or olfactory signals. Most flowers offer pollinators a reward (e.g. nectar and pollen) for the pollination service. However, one‐third of Orchidaceae have been shown not to provide a reward. Calanthe are terrestrial orchids distributed throughout China, Nepal, Japan and tropical Asia. Despite its high diversity, the pollination biology of Calanthe remains largely unknown, even though it is an important aspect of plant conservation. In the study, through field surveying, there were three Hesperiidae butterflies pollinating two species of Calanthe and the pollination behavior differed between the two species of Calanthe, which might lead to different fruit setting rates. There was no nectar in the flowers of the two species, indicating deceptive pollination. Using a glass cylinder experiment, it was deduced that the two species of Calanthe were most likely to attract pollinators by generalized food deception. Interestingly, Hesperiidae butterflies were traditionally thought to be nectar thieves and generally do not transmit pollinia. However, our findings showed that, in this case, the thieves were deceived by the plants and pollinated them for free.     

Effect of nectar supplementation on male and female components of pollination success in the deceptive orchid Dactylorhiza sambucina

Many orchids lack floral nectar rewards and therefore rely on deception to attract pollinators. To determine the effect that a mutation for nectar production would have on overall pollination success of the deceptive orchid Dactylorhiza sambucina, we recorded pollen deposition and removal in flowers of plants that had either been supplemented with an artificial nectar solution or left unmanipulated as controls. Nectar supplementation resulted in significant increases in the proportion of flowers pollinated, regardless of morph colour and the density of plants supplemented in the population. However, nectar supplementation had a significant positive effect on pollinaria removal only for the yellow morph in one experiment in which a low proportion of plants were supplemented. Thus a mutation for nectar production would have a positive effect on overall pollination success in D. sambucina, particularly the female component. The observed patterns are discussed in relation to other factors, such as cross-pollination and the reallocation of nectar resources for other plant functions, which are traditionally considered to shape the rewardless strategies of orchids.     

The dynamics of pollen removal and deposition,and its effects on sexual phases in a protandrous plant: Glechoma longituba (Lamiaceae)

The duration of sexual phases in dichogamous plants are affected by many factors. Using both experimental and observational studies, we investigated natural patterns of pollen removal and deposition, visiting frequency of pollinators, patterns of nectar secretion, and effects of pollen removal and stigmatic pollen deposition on the duration of sexual phases in a protandrous plant, Glechoma longituba. We found that visiting frequency of pollinators correlated with the nectar secretion pattern. The nectar volume during the male phase was higher than during the female phase. In the morning, the main pollinator, the bee Anthophora plumipes, mainly foraged for nectar and showed no preference for flowers in male or female phase, despite male phase flowers producing higher amounts of nectar. However, in the afternoon, they changed their behavior and foraged mainly for pollen, and then showed a preference for flowers in male phase. Furthermore, the rates of pollen removal and stigmatic pollen deposition can affect the starting time and the duration of the female phase. When pollen removal and pollination rates are low due to scarcity of pollinator services, the sexual phase can be prolonged, leading to an overlap, and thereby enhance the chance for sexual reproduction through pollinator‐facilitated self‐pollination. We consider the variation of sexual phases in Glechoma longituba an adaptive mechanism prepared for both cross‐pollination enhancement and reproductive assurance depending on the available pollination services.     

Ancestral deceit and labile evolution of nectar production in the African orchid genus Disa

An outstanding feature of the orchid family is that approximately 30–40% of the species have non-rewarding flowers and deploy various modes of deception to attract pollinators, whereas the remaining species engage in pollination mutualisms based on provision of floral rewards. Here, we explore the direction, frequency and reversibility of transitions between deceptive and rewarding pollination systems in the radiation of the large African genus Disa, and test whether these transitions had consequences for diversification. By optimizing nectar production data for 111 species on a well-resolved phylogeny, we confirmed that floral deception was the ancestral condition and that nectar production evolved at least nine times and was lost at least once. Transitions to nectar production first occurred ca 17 million years ago but did not significantly affect either speciation or extinction rates. Nectar evolved independently of a spur, which was lost and gained multiple times. These results show that nectar production can be a highly labile trait and highlight the need for further studies of the genetic architecture of nectar production and the selective factors underlying transitions between deception and mutualism.     

新疆荒漠植物耳叶补血草的爆发式开花式样与传粉模式

荒漠环境中干热、大风、浮尘、温度骤变等突发性天气状况,对植物的有性生殖具不利影响.耳叶补血草(Limonium otolepis)是新疆盐生荒漠的主要建群种,对其适应荒漠恶劣环境的生殖策略仍缺乏研究.为了探讨荒漠植物适应环境的开花式样和传粉模式,作者通过野外定点观测和室内电镜扫描等方法对耳叶补血草的开花进程、花部特征、传粉媒介、结实特性等进行了研究.耳叶补血草的花期在6月上旬至7月中旬,花期较长且具明显不同步的单株开花状态.在一天中,单花表现出极为集中的开花、散粉过程:一般早上8:00开始开放,9:00达开放高峰期,11:00以后基本不再有单花开放;开放后的单花持续6-8 h后闭合.每单花平均有花粉752粒,花粉表面有网状纹饰,有花蜜.传粉者主要为蜜蜂科和食蚜蝇科昆虫,访花高峰期在9:00-14:00.人工授粉实验表明耳叶补血草以异交为主,自动自花授粉率低.自然状态下结实率为36％,在花序中基部第一位上单花的结实率(45.7％)明显高于第二位上单花(3.3％).耳叶补血草爆发式开花、在短时间内快速完成传粉过程,可能是躲避荒漠环境突发性天气条件的一种策略;而居群分散的开花时间及较长的花期在应对不稳定生境,分摊生殖风险方面具重要意义.     

Pollination and breeding system of Couepia uiti (Mart. and Zucc.) Benth (Chrysobalanaceae) in the Pantanal da Nhecolandia.

The pollination biology and breeding system of Couepia uiti was studied. In this species, flowers opened at 06:00 AM anthesis, and nectar production began at around 0800 h, reached a maximum volume from 09:30 AM to 10:30 AM, and decreased thereafter. The nectar sugar concentration increased continuously, but showed an abrupt increase from 10:00 AM to 12:00 AM. Pollen release occurred at about 09:30 AM and was quickly collected. The stigmas became receptive at around 12:00 AM. The pollinators of C. uiti included the bees Apis mellifera, Xylocopa sp. and Bombus sp., and three species of wasps. This conclusion was based on the observation that these hymenopterans had C. uiti pollen on their bodies, visited the receptive flowers, and touched the anthers and stigmas, thereby promoting pollination. Of these floral visitors, A. mellifera was considered to be the most efficient pollinator. However, mixed pollination also occurred. The number of C. uiti flowers visited in the morning (n = 52) was three times smaller than in the afternoon (n = 62), and the species richness of floral visitors was also bigger in the afternoon (eight in the afternoon versus five in the morning). This finding indicated that these floral visitors preferred to exploit nectar rather than pollen. Controlled pollination experiments showed that C. uiti was a self-incompatible species that produced fruits only by cross-pollination. Treatments such as agamospermy and spontaneous and self-pollinations did not produce fruits.     

An Examination of the Function of Male Flowers in an Andromonoecious Shrub Capparis spinosa

The pollen donor and pollinator attractor hypotheses are explanations for the functions of the male flowers of andromonoecious plants. We tested these two hypotheses in the andromonoecious shrub Capparis spinosa L. (Capparaceae) and confirmed that pollen production and cumulative volume and sugar concentration of nectar do not differ between male and perfect flowers. However, male flowers produced larger anthers, larger pollen grains and smaller ovaries than perfect flowers. Observations on pollinators indicated that two major pollinators (Xylocopa valga Gerst and Proxylocopa sinensis Wu) did not discriminate between flower morphs and that they transferred pollen grains a similar distance. However, there were more seeds per fruit following hand pollination with pollen from male flowers than from perfect flowers. Individuals of C. spinosa with a larger floral display (i.e. bearing more flowers) received more pollen grains on the stigma of perfect flowers. Female reproductive success probably is not limited by pollen. These results indicate that male flowers of C. spinosa save resources for female function and that they primarily serve to attract pollinators as pollen donors.     

兰科植物欺骗性传粉

植物与传粉动物的互利关系在生态系统中非常普遍。然而,有许多植物不为传粉者提供任何报酬,而是利用各种欺骗方式诱骗昆虫拜访,从而实现传粉,即欺骗性传粉。兰科是被子植物大科之一,其高度特化的繁殖器官和适应于昆虫传粉的精巧结构令人称奇。进化论创始人达尔文描述了许多兰花与昆虫精巧的传粉系统,但他忽视了欺骗性传粉的存在。事实上,近1/3的兰科植物都依赖于欺骗性传粉。欺骗性传粉可能是导致兰科植物多样性的重要原因之一。兰花利用或操作昆虫觅食、交配、产卵和栖息等行为,演化出各种各样的欺骗性传粉机制,常见的类型包括泛化的食源性欺骗、Batesian拟态、性欺骗、产卵地拟态和栖息地拟态。花的颜色、形态和气味在欺骗性传粉的成功实现中起到了重要作用。欺骗性兰花与传粉昆虫之间的演化可能是不同步的,兰花追踪昆虫的行为信号而发生分化,然而欺骗性传粉可能对昆虫造成一定的伤害,从而对昆虫也施加选择压力。由于昆虫的学习行为,欺骗性的兰花一般具有低的昆虫拜访率和结实率,其繁殖成功率受各种因素的影响。欺骗性加剧了兰花对传粉昆虫的依赖,使其具有更高的灭绝风险,传粉生物学的研究能为兰科植物的有效保护提供指导。在欺骗性传粉系统中,有报酬的伴生植物、拟态模型和其他拟态信号提供者对传粉成功有重要影响。因此,研究欺骗性传粉兰花、传粉昆虫和相关的生物和生态因子的网状进化关系具有重要理论和实践意义。     

Pollen in Bee-Flower Relations Some Considerations on Melittophily*

Pollen and nectar are usually lumped together as floral rewards for pollinating bees, but they play totally different roles for flowers and bees (Table 1), as well as in the relationship between them. While flowers are specialized for certain pollinators via nectar, bees specialize on certain flowers via pollen. While flowers need pollen as a prerequisite for pollination, it is the essential larval food in bees. Thus, there is a strong competition between them for pollen. Foraging for pollen must be divided into three phases: uptake in the flower, reloading into and homeward transport within a carrying container. Bees have specializations for transport but hardly any for pollen uptake - and thus for pollination. Bees actively harvesting pollen usually do not pollinate. This only happens as a consequence of contamination of the bee by pollen. From these data a scenario is provided for the evolution of bees and bee flowers. Specialized bee flowers are often characterized by their ability to hide pollen from the bees and at the same time use them as optimal pollinators. If the relationship of bees and flowers is mutualistic at all it is best described as a balanced mutual exploitation.     

Mechanisms and evolution of deceptive pollination in orchids

The orchid family is renowned for its enormous diversity of pollination mechanisms and unusually high occurrence of non-rewarding flowers compared to other plant families. The mechanisms of deception in orchids include generalized food deception, food-deceptive floral mimicry, brood-site imitation, shelter imitation, pseudoantagonism, rendezvous attraction and sexual deception. Generalized food deception is the most common mechanism (reported in 38 genera) followed by sexual deception (18 genera). Floral deception in orchids has been intensively studied since Darwin, but the evolution of non-rewarding flowers still presents a major puzzle for evolutionary biology. The two principal hypotheses as to how deception could increase fitness in plants are (i) reallocation of resources associated with reward production to flowering and seed production, and (ii) higher levels of cross-pollination due to pollinators visiting fewer flowers on non-rewarding plants, resulting in more outcrossed progeny and more efficient pollen export. Biologists have also tried to explain why deception is overrepresented in the orchid family. These explanations include: (i) efficient removal and deposition of pollinaria from orchid flowers in a single pollinator visit, thus obviating the need for rewards to entice multiple visits from pollinators; (ii) efficient transport of orchid pollen, thus requiring less reward-induced pollinator constancy; (iii) low-density populations in many orchids, thus limiting the learning of associations of floral phenotypes and rewards by pollinators; (iv) packaging of pollen in pollinaria with limited carry-over from flower to flower, thus increasing the risks of geitonogamous self-pollination when pollinators visit many flowers on rewarding plants. All of these general and orchid-specific hypotheses are difficult to reconcile with the well-established pattern for rewardlessness to result in low pollinator visitation rates and consequently low levels of fruit production. Arguments that deception evolves because rewards are costly are particularly problematic in that small amounts of nectar are unlikely to have a significant effect on the energy budget of orchids, and because reproduction in orchids is often severely pollen-, rather than resource-limited. Several recent experimental studies have shown that deception promotes cross-pollination, but it remains unknown whether actual outcrossing rates are generally higher in deceptive orchids. Our review of the literature shows that there is currently no evidence that deceptive orchids carry higher levels of genetic load (an indirect measure of outcrossing rate) than their rewarding counterparts. Cross-pollination does, however, result in dramatic increases in seed quality in almost all orchids and has the potential to increase pollen export (by reducing pollen discounting). We suggest that floral deception is particularly beneficial, because of its promotion of outcrossing, when pollinators are abundant, but that when pollinators are consistently rare, selection may favour a nectar reward or a shift to autopollination. Given that nectar-rewardlessness is likely to have been the ancestral condition in orchids and yet is evolutionarily labile, more attention will need to be given to explanations as to why deception constitutes an 'evolutionarily stable strategy'.