POLLINATION ADAPTATION IN PEDICULARIS LANCEOLATA

Cinematographic, stereophotographic, and visual observations have revealed that the zygomorphic, nectarless flower of Pedicularis lanceolata Michx. is pollinated in northeastern Ohio by pollen-foraging workers of three Bombus species. Foragers hang inverted from the asymmetrical corolla and remove pollen with their front and middle legs from anthers concealed by the galea. The stigma contacts pollen deposited especially on the median portion of the forager's first abdominal sterilite. Insect exclosures and pollinator collections demonstrated the obligate pollination dependence of the flower on Bombus workers. Of 152 corbicular pollen loads from workers foraging on P. lanceolata, Gentiana andrewsii, Prunella vulgaris and five composite species in the same habitat, 84.2% contained P. lanceolata pollen; 75.8% of the latter contained P. lanceolata pollen exclusively. Possible coadaptive relationships of Pedicularis flowers and pollinators are discussed.     

POLLINATION ADAPTATION IN PEDICULARIS CANADENSIS

During the spring of 1966 and 1967, operation of the pollination mechanism of Pedicularis canadensis L. flowers by nectar- and pollen-foraging bumblebee (Bombus Latr.) queens was observed in an Iowa relict prairie and was recorded by high-speed cinematography. The stigma and style were found to sweep the forager's pronotum crevice as head and tongue were introduced into the 12-mm corolla tube. Pollen deposited in the crevice was not removed as the forager's middle leg only groomed pollen laterally deposited on head and scutum. Thirty corbicular pollen loads from 417 foragers collected on nine prairie plant species included pollen from Pedicularis canadensis, Aquilegia canadensis L., Dodecatheon meadia L., and two unidentified plants not found in the prairie. The pollination mechanism was considered closely adapted morphologically and phenologically to bumblebee queens of eight species found foraging on Pedicularis.     

THE POLLINATION ECOLOGY OF PEDICULARIS ON MOUNT RAINIER

Six native species of Pedicularis in Mount Rainier National Park in Washington State were studied for their reproductive relationships with animal pollinators. Cinematographic and stereophotographic records revealed pollination of the nectariferous P. bracteosa and P. rainierensis by upright, nectar-foraging queens and workers of five bumblebee (Bombus Latr.) species and by inverted workers scraping pollen from anthers concealed within the corolla galea. The nectarless, rostrate flowers of P. contorta, P. groenlandica, P. ornithorhyncha, and P. racemosa were pollinated by pollen-foraging workers and occasional queens virbrating pollen from concealed anthers. Insect exclosure methods revealed complete absence of fruiting in the absence of insects, and pollinator collections further indicate obligate dependence of the plants upon bumblebees for their sexual reproduction. Analysis of corbicular pollen loads from pollinators suggested that pollinator species are not monolectic but that individual pollinators range from monolectic to polylectic. Measurements indicated limited correlations between lengths of corolla tubes and tongues of nectar-foraging insects. Each nectarless Pedicularis species occupied a different, specific habitat, but P. bracteosa and the endemic P. rainierensis were sympatric in part. Each species had a unique spectral reflectance pattern from the corolla. Proximity of habitats and overlap of blooming periods of all Pedicularis species eliminate the possibility of contemporary geographic or phenological reproductive isolation. It is suggested that behavioral interactions of the plants and their insect pollinators may have been instrumental in the past in reproductively isolating these species, hybrids of which are unknown.     

滇西北马先蒿属传粉生物学的初步研究

对云南西北部的中甸和德钦高山地区４种马先蒿属（ＰｅｄｉｃｕｌａｒｉｓＬ．）植物的传粉生物学进行了观察。晚夏开花的马先蒿与传粉熊蜂（ＢｏｍｂｕｓＬａｔｒ．）的出现在物候上有一定的相关性。其中短管、无喙、无蜜腺的肉根马先蒿（Ｐ．ｒｏｙｌｅｉＭａｘｉｍ．）和茨口马先蒿（Ｐ．ｔｓｅｋｏｕｅｎｓｉｓＢｏｎａｔｉ）是由熊蜂工蜂震动花粉、背触式传粉,这与其花冠类型的较原始性有关;长管、具喙、无蜜腺的斑唇马先蒿（Ｐ．ｌｏｎｇｉｆｌｏｒａｖａｒ．ｔｕｂｉｆｏｒｍｉｓ（Ｋｌｏｔｚ．）Ｔｓｏｏｎｇ）和云南长管马先蒿（Ｐ．ｓｉｐｈｏｎａｎｔｈａＤｏｎ．ｖａｒ．ｄｅｌａｖａｙｉ（Ｆｒａｎｃｈ．）Ｔｓｏｏｎｇ）通过熊蜂工蜂震动花粉、腹触式传粉。长管类马先蒿在植株相对矮小的情况下,通过较长的花冠管具喙向外伸出以实现授粉,这可能是与其传粉熊蜂协同进化的结果。     

THE POLLINATION DYNAMICS OF SYMPATRIC SPECIES OF PEDICULARIS (SCROPHULARIACEAE)

Sympatric populations of Pedicularis oederi, P. cystopteridifolia, and P. groenlandica on the Beartooth Plateau (Montana) were obligately dependent on Bombus pollinators. Their corolla colors were mutually distinct to insect vision, but their nectars had identical sugar components. Analysis of corbicular pollen loads of pollinating insects indicated a high degree of polylecty. Queen/worker ratios of pollinators on plant species corresponded to the parallel phenological sequences of plant blooming and insect caste development. The total number of individual pollinators of each Bombus species on all plants was directly related to the number of plant species it pollinated, but a comparable relationship between the number of pollinators on a plant species to the number of Bombus species pollinating it was not found. Morphological and behavioral correspondence of floral mechanisms and pollinators form an integral part of the general pattern of coadaptive evolution of the floral ecology of Pedicularis in North America known from previous studies. Reproductive isolation between Pedicularis species in this study is attributed to internal barriers. Wide overlaps of blooming seasons, proboscis lengths of bumblebee species and castes, and pollinator species on Pedicularis species suggest resource sharing, rather than resource partitioning or competition for resources among plant and insect species and individuals.     

RATES OF FLORAL EVOLUTION: ADAPTATION TO BUMBLEBEE POLLINATION IN AN ALPINE WILDFLOWER,POLEMONIUM VISCOSUM

Animal pollinators are thought to shape floral evolution, yet the tempo of this process has seldom been measured. I used the prediction equation of quantitative genetics, R = h²S , to predict the rate at which a change in pollinator abundance may have caused divergence in floral morphology of the alpine skypilot, Polemonium viscosum. A selection experiment determined the rate at which such divergence can actually proceed. Corolla flare in this species increases by 12% from populations pollinated by a wide assemblage of insect visitors to those pollinated only by bumblebees. To simulate the evolutionary process giving rise to this change, I used a pollinator selection experiment. Plants with broad flowers set significantly more seeds than plants with narrow flowers under bumblebee pollination but had equivalent fecundity when visited by other insects or hand-pollinated. Bumblebee-mediated selection for broad corolla flare intensified from 0.07 at seed set to 0.17 at progeny establishment. Maternal parent-offspring regression yielded a confidence interval of 0.22–1.00 for trait heritability. Given these parameter estimates, the prediction equation shows that broadly flared flowers of bumblebee-pollinated P. viscosum could have evolved from narrower ones in a single generation. This prediction is matched by an observed 9% increase in offspring corolla flare after a single bout of bumblebee-mediated selection, relative to offspring of unselected controls. Findings show that plant populations can adapt rapidly to abrupt changes in pollinator assemblages.     

ASSORTATIVE POLLINATION IN LYTHRUM

The pollination patterns of bees and butterflies were observed in an assemblage composed of 57 plants of Lythrum alatum and 30 plants of L. salicaria. Lythrum salicaria was preferred by both groups of pollinators as indicated by the greater number of flights from this species as compared to flights from L. alatum. The degree of assortative pollination was measured in both groups of pollinators and was greater in bees. Fidelity in bees appeared to be in response to species attractiveness, whereas in butterflies it appeared to be in response to species balance.     

DARK ADAPTATION IN DINEUTES

The level of dark adaptation of the whirligig beetle can be measured in terms of the threshold intensity calling forth a response. The course of dark adaptation was determined at levels of light adaptation of 6.5, 91.6, and 6100 foot-candles. All data can be fitted by the same curve. This indicates that dark adaptation follows parts of the same course irrespective of the level of light adaptation. The intensity of the adapting light determines the level at which dark adaptation will begin. The relation between log aI ₀ (instantaneous threshold) and log of adapting light intensity is linear over the range studied.     

DARK ADAPTATION IN AGRIOLIMAX

A method is described which measures the excitation of Agriolimax by light, during the progress of light adaptation, by assuming that the orientating effect of continuous excitation is expressed as a directly proportionate tension difference in the orienting muscles of the two sides of the body. The tendency toward establishment of such a tension difference is caused to work against a similar geotropic effect at right angles to the phototropic one. This enables one to study the kinetics of light adaptation, and of dark adaptation as well. The situation in the receptors is adequately described by the paradigm See PDF for Equation similar to that derived by Hecht for the differential sensitivity of various forms, but with the difference that the "dark" reaction is not only "bimolecular" but also autocatalysed by the reaction product S. The progress of dark adaptation is reflected (1) in the recovery of the amplitude of the orientation and (2) in the rates of light adaptation at different levels of the recovery; each independently supports these assumptions, for which the necessary equations have been provided. These equations also account for the relative variabilities of the angles of orientation, and, more significantly, for the two quite different kinds of curves of dark adaptation which are obtained in slightly different types of tests.     

POLLINATION INTERACTIONS IN SYMPATRIC DICENTRA SPECIES

The pollination of a large mixed population of Dicentra canadensis and D. cucullaria was studied in a beech-maple forest in southwestern Ohio. Analyses of pollinator frequencies and species, pollinator behavior, sugar components and concentrations in nectar, corbicular pollen loads carried by pollinators, spectral reflectance from corollas including ultraviolet reflectance, fragrance, blooming phenology, insect-related fertility, and hybridization potentials suggested that these sympatric Dicentra species are reproductively isolated by internal mechanisms but not by external barriers to hybridization. Both species were exclusively and obligately dependent upon queen bumblebees for their pollination unlike other members of the spring ephemeral flora to which they belong. The occurrence of one or both species in one area is considered primarily the result of their adaptive preference for different edaphic and possibly climatic situations and not the consequence of competitive exclusion by pollinator behavior.     

ADAPTATION IN PANTOTHENATE-REQUIRING NEUROSPORA II. NUCLEAR COMPETITION DURING ADAPTATION

Davis , R. H. (U. Michigan, Ann Arbor.) Adaptation in pantothenate-requiring Neurospora. II. Nuclear competition during adaptation. Amer. Jour. Bot. 47(8) : 648–654. Illus. 1960.—The process of adaptation in pantothenate-requiring Neurospora was studied by the use of heterocaryons constituted of the pan-1 strain and a modified strain, pan-1 m, derived from it. Although pan-1-m homocaryons grow well on a pantothenate concentration on which pan-1 grows little or not at all, pan-1 nuclei often have a selective advantage in pan-1 + pan-1-m heterocaryons grown on the same medium. This results in non-adaptive changes in nuclear ratios and labile growth rates of the heterocaryons. Nuclear competition does not occur when pantothenate is not limiting to the growth of either homocaryon. The results are discussed, and related to past work on adaptation and nuclear ratio changes in Neurospora hetercaryons.     

PHOCINE DISTEMPER VIRUS INFECTION IN HARP SEALS (PHOCA GROENLANDICA)

Antibodies against phocine distemper virus were found in liver and serum specimens of harp seals from the Barents Sea and north of Jan Mayen (West Ice) in 1987 and 1989. By immunoperoxidase testing, antibodies against phocine distemper virus could be demonstrated in 34 of 74 liver specimens (46%) from 1987 and in 44 of 126 liver specimens (35%) from 1989. The prevalence of neutralizing serum antibodies against phocine distemper virus was found to be 98% in 1989. Both findings clearly demonstrate the presence of a widespread infection with phocine distemper virus or a closely related virus in harp seals. The significance of these findings to the recent outbreak of phocine distemper virus infection in harbor seals in Northwestern Europe is discussed and it is concluded that there may be an epizootiological link between this outbreak and the large migration of harp seals in 1987.     

EFFECTS OF POLLINATION INTENSITY IN CAMPSIS RADICANS

I examined effects of pollination intensity on fruit, seed and seedling characteristics in Campsis radicans, and joint effects of pollen donor and pollination intensity on fruit production. Large pollen loads were more likely to initiate fruit production than small pollen loads, and the former fruits contained more seeds and a greater total seed mass. No further increases in seed number or mass occurred for pollen loads above 4,000 grains. The weight of individual seeds was unaffected by pollen load. Effects of pollen donor were generally larger than effects of pollen load, and fruit production from small loads of pollen from one donor were sometimes equal to fruit production from larger pollen loads from another donor. The ratio of pollen grains deposited to resultant seeds increased with pollen load, and several explanations are proposed. Seeds from heavy pollinations emerged better than seeds from light pollinations, but did not differ in speed of germination or in the performance of seedlings up to 126 days. The emergence differences are probably due to differing intensities of pollen tube competition.     

POLLINATION AND POLLEN ENERGETICS IN MEDITERRANEAN ECOSYSTEMS

Most plants in Mediterranean ecosystems are insect pollinated, with pollen being the main reward to pollinators. The great majority of pollinators (70%) are bees and flies. We measured the energy content of pollen from 40 plant species in these ecosystems that represent abiotic and biotic pollination modes as well as the number of species of their pollinators. Pollen energy content correlates with pollinator diversity. Pollen of wind-pollinated plants contained less energy than that of insect-pollinated plants; there was no difference between insect-pollinated dicots and insect-pollinated monocots. The median date of flowering (from 1 January) estimated for each of the plant species did not vary significantly either with the number of pollinator species or with the energy content of pollen. The reasons for the differing values of pollen energy content are discussed; in particular, its relation to the type of pollen reserves, length of the flower style, and pollination enhancement.