草乌传粉过程中的广告效应与回报物质研究

虫媒传粉植物与其传粉者之间的相互作用被认为是被子植物花多样性的一个重要选择压力。这种相互作用体现在植物对传粉者的吸引以及传粉者行为对花粉的转运两个方面。本项研究通过去除不同的花部器官研究了草乌(Aconitum kusnezoffii)对其传粉者的吸引, 并结合传粉者的访问行为和草乌花的生物学特性探讨了传粉过程与交配系统的适应。红光熊蜂(Bombus ignites)是草乌的有效传粉者。去除花萼片显著降低了红光熊蜂的访问频率, 但去除特化成蜜腺叶的花瓣并没有显著改变红光熊蜂的访问频率, 这表明草乌吸引红光熊蜂的主要结构是由5枚萼片组成的花部外观形态, 而非花瓣。花蜜是草乌提供给红光熊蜂的回报物质, 糖浓度为39.23%, 组氨酸浓度为0.25 μg/μL。草乌花较大、单花花期长、雌雄异熟、花粉寿命长, 是一个自交亲和但需传粉者传粉完成繁殖过程的异交物种。草乌花序是无限花序, 当上部的花处于雄性阶段时, 下部的花正好处于雌性阶段。而红光熊蜂在草乌花序上的访问顺序通常自下而上, 带来异交花粉为下部的花进行异花授精, 同时又带走上部花的花粉, 这就很好地促进了草乌的异交。草乌雄蕊自外向内逐渐成熟, 是一种有效的限制传粉者单次访问浪费大量花粉的花粉装配策略, 能提高植物通过花粉散布获得的雄性适合度。     

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。     

露蕊乌头(毛茛科)不同海拔居群的传粉生态学

对青藏高原东北部两个不同海拔地点（互助,2460m和海北,3200m）的露蕊乌头Aconitum gymnandrum的传粉生态学进行了比较研究。研究结果表明高海拔居群的单花花期、雄性期持续时间、雌性期持续时间比低海拔长。露蕊乌头不存在无融合生殖现象,尽管高度自交亲和,但产生种子必须依赖传粉媒介。熊蜂是露蕊乌头的主要传粉昆虫,两个居群均存在传粉限制。熊蜂取食花蜜为主,在同一个花序上的访问顺序以由下向上为主,但在互助居群和海北居群有3．9％和2．7％的访花是由上向下进行的。统计发现有37．7％和29．3％的访问行为发生在同一植株内;因此,同株异花传粉造成的自交在露蕊乌头中仍然不能避免。低海拔居群的熊蜂种类和访花频率均高于高海拔居群,但自然状态下两个居群的结实率并没有显著差异。高海拔露蕊乌头居群可能具有较长的柱头持续时间,从而补偿了由于访花频率低带来的结实率降低。     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

管花秦艽的传粉生态学研究——兼与同域分布近缘种的比较

在青藏高原东北部连续两年观察了晚秋开花植物管花秦艽Gentiana siphonantha的传粉生态学特征,并在此基础上进一步比较分析了与该物种同域分布且亲缘关系较近、但开花较早的麻花艽G straminea之间的传粉生态学特征.管花秦艽的花发育过程表现出雌雄异熟和雌雄异位的特点,不存在花内的自花传粉,套袋隔离的花不结实也支持这一结论;株内自交的高结实率表明该物种是自交亲和的.盛花期每植株平均有15朵开放的花,雄性和雌性阶段的花比例为1.2:1;自然条件下产生种子必须依赖传粉媒介;苏氏熊蜂是最有效的传粉昆虫,且访花过程中埘雄性和雌性阶段花不具明显的偏向性;株内连续访花的频率高达87.8%,从而导致同株异花传粉自交的广泛存在.与同域分布的麻花艽相比,管花秦艽的单花花期、雄性和雌性期持续时间缩短.但盛花期开花数量明显增加.令人感兴趣的是尽管两个近缘种的花形态特征存在显著差异,但都是由同一种熊蜂传粉.这一特点与过去认为花颜色和花管长度是物种分化过程中与不同传粉昆虫协同进化导致牛殖隔离的假说不相符合.管花秦艽单花的访花频率和同株异花连续访花的比例都明显高于麻花艽.两个物种不同花序设计导致访花昆虫行为的改变可能是造成这一差异的主要原因.两个物种具有不同的开花时间,但仍然存在一定的花期重叠,表现出不完全的传粉生殖隔离状态.     

西方蜜蜂和兰州熊蜂在设施桃园的访花偏好性比较

【目的】为了比较西方蜜蜂 Apis mellifera 和兰州熊蜂 Bombus lantschouensis 在设施桃园内对不同时期桃花的访花偏好性、以及这种偏好性与花粉活力和采集花粉花蜜之间的关系。【方法】记录2种蜂在温室桃园内访问早期花、中期花和晚期花的比例,测定桃花不同时期的花粉活力以及2种蜂携带的花粉活力,观察2种蜂采集花蜜和采集花粉的成功率,统计2种蜂访花过程中桃花所处的枝条数及植株数。【结果】桃花不同时期的花粉活力差异显著,早期花花药未开裂,花粉未释放,中期花花粉活力（58.3%）显著高于晚期花花粉活力（34.2%）（P<0.01）;西方蜜蜂更加偏好访问中期花,对中期花的访问率高达75.3%,显著高于兰州熊蜂对中期花的访问率（49.2%）（P<0.01）;西方蜜蜂携带的花粉活力（92.1%）显著高于兰州熊蜂携带的花粉活力（72.9%）,但是西方蜜蜂采集花粉和采集花蜜的成功率均低于兰州熊蜂（P<0.01）;在访问一定数量的桃花过程中,兰州熊蜂在设施桃园内访问的枝条数和植株数较多,分布范围较广（P<0.01）。【结论】和兰州熊蜂相比,西方蜜蜂对活力花粉的辨别能力更强,更加偏好访问花粉活力较高的花朵,这种偏好性导致其采集花粉花蜜的成功率降低。     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

3种雌雄异熟樟科植物开花动态的比较研究

为了解樟科植物雌雄异熟的繁育系统特点,对3种樟科植物阴香(Cinnamomum burmannii)、紫楠(Phoebe sheareri)和浙江楠(Phoebe chekiangensis)雌雄异熟花的开花动态进行了比较研究。结果表明,3种植物雌性功能期的开始时间、雌性功能期和雄性功能期的时间分配有差异。3种植物的主要传粉者均为中华蜜蜂,在雌性功能期和雄性功能期均有访花;在雌性功能期第四轮退化雄蕊蜜腺和雄性功能期第三轮花丝基部的蜜腺分泌花蜜,不同开花阶段的花蜜蜜量和含糖量没有显著差异;柱头可授性和花粉活力在雌性和雄性功能期均较高;雌性功能期和雄性功能期均有自然花粉落置;座果率均是异交补粉自然套袋,表明都存在一定的传粉限制。因此,樟科植物雌、雄性阶段时间分配的不同,可能与退化雄蕊蜜腺和第三轮雄蕊附属蜜腺的分泌节律有关。     

华凤仙花部特征和传粉系统研究

研究了华凤仙（Impatiens chinensis L.）的花部特征和传粉系统,结果表明华凤仙单花期为2.3 d,雄性期是雌性期的6.8倍。花蜜量和花蜜可溶性糖含量分别为0.56 μL和28.2%。开花当天柱头即具有可授性,花粉活力在花朵刚开放时最高（95%）,至花朵凋谢之前仍保持较高的活力（55%）。花粉胚珠比（5 730.3±2 941.1）和套袋实验的结果表明华凤仙属于自交亲和但以异交为主的繁育系统,传粉过程需要传粉者。蜜蜂、熊蜂、蝶类和几种芦蜂是华凤仙的主要访花者,最有效的传粉者为三条熊蜂。考氏无垫蜂作为盗蜜者,可能是除正常访花者之外,对华凤仙的有性繁殖影响最大的生物因素。     

明亮熊蜂和意大利蜜蜂在温室桃园的访花行为和传粉生态学比较

2004—2006年连续3年应用明亮熊蜂和意大利蜜蜂在北京为温室桃园传粉,对其访花行为和传粉生态学进行研究.结果表明：两种蜂都可以替代人工掸花为温室桃园提供有效的传粉服务.两种蜂的访花行为和传粉效果不同,明亮熊蜂偏爱于采集花粉,主要以震动翅膀的方式来使花粉释放和传播,而意大利蜜蜂偏爱于采集花蜜,主要以身体接触的方式来粘附和传播花粉;明亮熊蜂的活动起点温度低、日工作时间长、访花速度快,在低温条件下比意大利蜜蜂的传粉效果好;意大利蜜蜂的趋光性强,飞撞温室塑料薄膜的现象严重,受温度和光照条件的影响较大,对温室环境的适应性较差.     

Remote perception of floral nectar by bumblebees

Summary On both artificial flowers in the laboratory and certain plant species in the field, bumblebees often closely approached flowers and then departed without probing for nectar. In laboratory experiments where nectar rewards were associated with subtle visual or olfactory cues, bumblebees approached and avoided non-rewarding flowers. Flowers that bees entered and probed for nectar contained rewards much more frequently than predicted by chance alone. When there were no external cues associated with nectar content, bees visited rewarding flowers by chance alone, provided rewarding flowers were not spatially clumped. In the field, bumblebees approached and rejected a large proportion of dogbane flowers and red clover inflorescences. On both species, flowers or inflorescences probed by bees contained more nectar than those rejected by bees or those that I chose at random. On fireweed and monkshood, bees rarely or never approached and rejected healthy-looking flowers. Predictions generated by an optimal foraging model were tested on data from four bumblebee species foraging on red clover. The model was highly successful in qualitatively predicting the relationship between handling time and proportion of inflorescences rejected by individual bees, and the relationship between threshold nectar content for acceptance by bees and average resource availability. Thus, bees appeared to use remotely perceived cues to maximize their rates of nectar intake.     

Optimal foraging in bumblebees and coevolution with their plants

Summary The aims of this paper were to consider the coevolution between bumblebee movement patterns within plants and various properties of the plants such as the spatial distribution of their flowers, and to determine the extent to which the bumblebees and the plants can be considered to be maximally adaptive or optimal. Attention was restricted to plants which have flowers arranged on vertical inflorescences and to the bumblebees which visit these plants.It was found that the bumblebees tend to commence foraging at the bottom of each infloresence, that they tend to move from one flower to the closest vertically higher flower, that they miss flowers as they move upwards and that they tend to leave each inflorescence before reaching the top. It was also found for the four common plant species considered that nectar abundance per flower decreases with flower height on an inflorescence, that the flowers with receptive stigmas are restricted to the bottoms of the inflorescences while the flowers shedding pollen occur above them, and that the flowers are arranged approximately in spirals on the inflorescences.The pattern of movements of the bumblebees and the various properties of the plants appear to represent coevolved adaptations. Furthermore the bumblebees' movement patterns appear to be optimal in the sense that they result in the maximum net rate of energy gain to the bumblebees. Further studies are necessary, however, to determine whether or not the plants can be considered to be optimal.An exception to the above scheme is provided by a plant which is quite uncommon in the study area. This plant also has flowers on vertical inflorescences and appears to be pollinated by bumblebees. However, while the pattern of movements of the bumblebees on this plant species are extremely similar to those on the four common species, this plant species exhibits quite different properties from the other four. Two possible explanations for this exception are presented.     

Contrasting movement patterns of nectar-collecting and pollen-collecting bumble bees (Bombus terricola) on fireweed (Chamaenerion angustifolium) inflorescences

Abstract. 1. Movements of nectar and pollen-foraging bumble bees on inflorescences of Chamaenerion angustifolium (L.) J. Holub (fireweed or rosebay willow herb) were compared with predictions based on reward distributions and optimality principles. Observations suggest that nectar and pollen-gathering bumble bees behave according to the same set of reward maximization criteria when foraging from flowers of this species.
2. Both kinds of foragers matched their arrival points with the vertical positions on inflorescences in which the densities of their respective food resources were greatest. For nectar-foragers, this point was located at the lowest tier of flowers, whereas for pollen-foragers it was found in the middle of the inflorescences. Nectar and pollen-foraging bees both moved upward on inflorescences following gradients from high to low reward availability.
3. Nectar-foragers responded to decreases in inflorescence size over the season by reducing the number of flower visits made on each raceme. Number of flowers visited by pollen-foragers was low throughout and reflected the scarcity of male-phase flowers on racemes. Flower revisitation rates were low for both kinds of workers, but were slightly higher for those collecting pollen.     

Geitonogamy in rewarding and unrewarding inflorescences: modelling pollen transfer on actual foraging sequences

Many orchid species are unusual in that they provide no nectar or pollen rewards for their pollinators. Absence of reward is expected to have a fundamental effect on pollinator visitation patterns. In particular the number of flowers visited per inflorescence is expected to be affected in both unrewarding and co-flowering rewarding species. We used arrays of artificial inflorescences, which could be either rewarding or unrewarding and were differentiated by their colour, to test how many flowers bumblebees visit in each type of inflorescence. The frequency of the two colours was varied, thus modelling the case where different frequencies of both an unrewarding and rewarding species were present in a patch. We found that bumblebees visited more flowers per rewarding inflorescence after they have experienced unrewarding or partially emptied rewarding inflorescences. We used these results to simulate pollen transfer and thus predict selfing rates on rewarding inflorescences. We found these increased when nectar depleted or when there was a greater proportion of unrewarding inflorescences in the patch. Conversely, we found that the number of flowers bumblebees visited on each unrewarding inflorescence did not significantly change through experiments. Selfing rates for unrewarding inflorescences were predicted to depend principally on the number of these inflorescences bumblebees visited rather than on the number of flowers they visit per inflorescence. This was because most visitors to orchids are supposed to be naive, and pollinators that commence foraging carrying no pollen will necessarily self any flower they pollinate on the first inflorescence they visit. Thus the average selfing rate is expected to increase as the sequence of inflorescences visited decreases in length.     

Pollen foraging by bumblebees: Foraging patterns and efficiency on Lupinus polyphyllus

Summary Bumblebees foraging on vertical inflorescences start near the bottom and work upward, behavior commonly interpreted as a response to the greater amounts of nectar available in lower flowers. Lupinus polyphyllus, which produces no nectar, has more pollen available in upper flowers. Although bees are probably unable to detect this gradient, since pollen is hidden from their view, they still start low and forage upward. Therefore, we concluded that the bees' tendency to forage upward on vertical inflorescences is not tied to a reward gradient. In addition, bees use only about 15% of the flowers per inflorescence, although they could be much more efficient by visiting and revisiting every flower systematically. In general, revisits would not be penalized because most flowers contain enough pollen for several visits. Optimal foraging theory may not offer an adequate explanation for such gross inefficiency.     

The foraging movements of bumblebees on vertical “inflorescences”: An experimental analysis

Summary The bumblebees,Bombus edwardsii, move upward while visiting consecutive flowers on artificial inflorescences. This response is unrelated to the vertical patterning of rewards in the flowers of inflorescences. However, when rewards are greatest in the bottommost flowers the bees learn to start lower and leave before reaching the topmost (empty) flowers. Conversely, when rewards are greatest in the topmost flowers they tend to start in the middle of the inflorescence and depart from the top. When rewards are equal in all flowers bees start near the bottom and depart near the top of inflorescences. These behavioral patterns tend to maximize the number of visits to rewarding flowers while minimizing visits to non-rewarding flowers, thereby enhancing foraging returns.Supported by NSF grant DEB77-08430.     

Multi-cycle synchronous protandry in raceme-like inflorescences of a bumblebee-pollinated herb Aconitum grossedentatum

Multi-cycle synchronous dichogamy is expected to be a mechanism for reducing self-pollination and sexual interference. It is often found in plants with umbellate inflorescences where pollinator movement is unpredictable, but not in plants with raceme inflorescences that are pollinated by bumblebees. Plants with raceme inflorescences often acropetally open flowers, resulting in an arrangement of females at lower level and males at upper level. This is good enough to preclude geitonogamy because bees tend to move upwardly within the inflorescences. Furthermore, although the degree of segregation of sexes varies among species, their intraspecific variations within a population have rarely been examined. Here, we present a synchronous protandry in bee-pollinated Aconitum grossedentatum, which has a raceme-like inflorescence and opens flowers basipetally. To evaluate the functional significance of synchronous dichogamy in mating, we firstly observed the distribution of sex phases of open flowers. Then, we assessed the effect of each phase flower on foraging behavior by pollinators and seed-set success. The inflorescences tended to exhibit either male- or female-phase flowers at any moment early in the flowering season, but the degree of segregation of sexes declined over time within a population. The degree of the segregation did not affect bumblebee visits to flowers, but it decreased seed-set success of female-phase flowers at that time. Our results demonstrated that synchronous protandry was beneficial for pollination success in A. grossedentatum by avoiding geitonogamy. Nevertheless, we also found asynchronous protandry late in the season, suggesting that the benefits by synchronous protandry decreased over the season.

     

The movement patterns of carpenter beesXylocopa micans and bumblebeesBombus pennsylvanicus onPontederia cordata inflorescences

The movement patterns of carpenter bees (Xylocopa micans) and bumblebees (Bombus pennsylvanicus) foraging for nectar on vertical inflorescences ofPontederia cordata were studied near Miami, Florida. The floral biology ofP. cordata is unique in several ways: (a) many short-lived flowers per inflorescence, (b) constant nectar production throughout the life span of each flower, and (c) abscence of vertical patterning of nectar and age of flowers. Inflorescences ranged between 3.5 and 15.8 cm long and had between 9 and 55 open flowers. Both carpenter bees and bumblebees arrived mostly on the bottom third of the inflorescence and left after visiting flowers on the top third of the inflorescence. The departure position from the inflorescence was higher up than observed in studies of other insect pollinators foraging on other speces of plants. This pattern of departure probably occurs in the absence of a vertical gradient of nectar or floral morphology.     

Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Why do nectar-foraging bees and wasps work upwards on inflorescences?

Summary Wasps (Dolichovespula and Vespula spp.) worked predominantly upwards when foraging for nectar on inflorescences of the protogynous Scrophularia aquatica, in which the standing crop of nectar sugar per flower showed no clear pattern of vertical distribution up an inflorescence. Bumblebees taking nectar (Bombus hortorum visiting legally, and certain individuals of B. terrestris which positioned themselves head-upwards while taking nectar through holes bitten in the corolla) worked predominantly upwards on the racemose inflorescences of Linaria vulgaris, although the standing crop of nectar sugar per open flower increased up the raceme. Individuals of B. terrestris which robbed Linaria flowers in a head-down position worked predominantly downwards on inflorescences. The upward or downward directionality of intra-inflorescence movements by foraging insects may depend in part on the position these adopt during their flower visits.