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South African Journal of Zoology

ISSN: 0254-1858 (Print) (Online) Journal homepage: http://www.tandfonline.com/loi/tafz19

Sodium and potassium concentrations in floral
nectars in relation to foraging by honey bees

S.W. Nicolson & P.V. W.-Worswick

To cite this article: S.W. Nicolson & P.V. W.-Worswick (1990) Sodium and potassium
concentrations in floral nectars in relation to foraging by honey bees, South African Journal of
Zoology, 25:2, 93-96, DOI: 10.1080/02541858.1990.11448196

To link to this article: http://dx.doi.org/10.1080/02541858.1990.11448196

Published online: 02 Oct 2015.

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the levels of sodium and potassium in the urine did not necessarily reflect those in the nectar being gathered. organic acids. Zoo!. to see whether low nectar sodium concentrations made the asssumption that the bees foraging on a parti- are a general phenomenon. Marondera.en kaliumkonsentrasies in nektar van verskeie spesies blomplante. except for Echium fastuosum. Nicolson* and P. lipids. haemolymph and cular plant were flower-constant for that species. may be ettes. Zimbabwe An important aspect of pollination biology concerns the robusta and four species of Myrtaceae. wat deur heuningbye (/V)is me/litera capensis) opgesoek word. the quantity of nectar was needs of flower visitors. and urine were then sampled using 1 or 2 Il-l micro- ber) from a variety of flowering plants upon which bees pipettes.). at various nator (reviewed by Heinrich 1975. 25(2) 93 Sodium and potassium concentrations in floral nectars in relation to foraging by honey bees S.3 mmol.Ll per inflorescence. The volume of nectar in each flower vitamins and inorganic minerals. alhoewel hemolimfkonsentrasies strang gereguleer word. Rondebosch. Hierdie resultate word vergelyk met die nektarioonkonsentrasies in blomme wat deur houtkapperbye en kolibries besoek word. Leucospermum).7 ± 4. techniques (Instrumentation Laboratory Model IL 243). amino acids. we forage. The haemolymph Nectar was collected during spring (September-Octo. the only systematic expressed as j. This is because sites including the campus of the University of Cape nectar is predominantly a sugar solution and most polli. South Africa. Feeny & Sodium and potassium concentrations of the nectar Lederhouse 1974.8 ± 1. Nicolson & Louw were measured by means of standard flame photometric 1982). then stored on ice and immediately transported Methods (10-15 min) back to the laboratory. Town and Kirstenbosch Botanic Gardens. Flower urine were examined in bees foraging on different nec. Natrium. en die gemiddelde kaliumkonsentrasie 18. Afr.en kaliumvlakke in die uriene nie noodwendig die van versamelde nektar weerspieel het nie. is bepaal.O. was determined from the column length in the micropip- ents of nectar.4 mmol (± S. as well as its water content. who examined a variety of using a portable refractometer (Bellingham and Stanley hummingbird flowers and found sodium concentrations Ltd).4 mmol (± S. and appropriate dilutions were made when either to be much lower than those of potassium. W. may be in short supply in their diet (Anns. In 18 plantspesies was die gemiddelde natriumkonsentrasie 9. Analysis of honey bee body fluids showed that. 2 or 10 Il-l) were used for & Baker (1975) emphasized that the nectar of flowering withdrawing nectar from the corolla: usually the visco- plants contains a complex array of chemicals in addition sity of the nectar was low enough for the micropipette to to sugars:.3 mmol. accepted 3 October 1989 Sodium and potassium concentrations have been measured in nectar from a variety of flowering plants visited by honey bees (/V)is me/litera capensis).-Worswick** Department of Zoology. These other constitu. Bees foraging on three chemistry of the body fluids. Box 49. proteins. 1983).S.). Baker (Drummond Scientific Co. However.8 ± 1.F. firstly. The attention. J.E. constancy in honey bees has been well documented tars. The aim of the present study was. Seeley 1985).7 ± 4.W. Barrows 1974. look at ion concentrations in floral nectars is the work of The sugar concentration of the nectar was measured Hiebert & Calder (1983). fill by capillarity. Where the flower consisted of an inflorescence important in filling the nutritional and homeostatic (e. to see whether nectar composition is reflected in the (Heinrich 1975. Flowers were energetics of the relationship between flower and polli. but there have been indications that sodium refractometer readings were converted to grams solute per 100 ml solution. P. Secondly. and the mean potassium concentration was 18. University of Cape Town.g. AU the species examined were indigenous to being obtained by gentle squeezing of the abdomen. However. These results are compared with nectar ion concentrations in carpenter bee and hummingbird flowers. 7700 Republic of South Africa Received 3 August 1989. the haemolymph being withdrawn through an (Apis mellifera capensis Escholtz) were observed to be incision in the third abdominal segment arid the urine foraging. • To whom correspondence should be addressed •• Present address: Donnervale Estate. 'n Analise van heuningbyliggaamsvloeistowwe het getoon dat. although haemolymph ion concentrations were closely regulated. The ionic the volume of nectar was too small or the sugar concen- balance of nectivorous insects has also received little tration exceeded the scale of the refractometer. to sample a In order to examine the ionic balance of honey bees selection of the flowering plants upon which honey bees foraging on nectars of varying ion concentrations.V. die natrium. different species were collected and killed with ethyl acetate. In 18 plant species the mean sodium concentration was 9. Grevillea Sodium and potassium concentrations of the body fluids . sampled in the early morning (8h00-10h00).. Micropipettes nators have very high energy demands. 1990.

3 Liliaceae Aloe ferox 12.4 mmol in Callistemon citrinus.001). cordifolium 30.5 10.3 13.2 12.E.7 15.6 ± 1.0 ± 0.7 ± 0.8 n = 6 for volume. Potassium con.1 14. Favre.2 5.2 ± 0. Lanum. with occasional visits to Leucospermum (Proteaceae).0 ± 2.5 ± 0.6 ± 0.0 ± 1.3 16.8 21.4 mmol and were not significantly sucrose equivalent.3 7.0 ± 0.3 L.0 ± 0.0 ± 0.5 ± 2. were significantly determining the energy content of these nectars directly lower than in the bee flowers sampled in the present from refractive index (Inouye.4 L. study (t test.3 13.5 C.5 ± 0. 25(2} were determined as for nectar.5 21. with a mean value of9.0 ± 0. Tydskr. Levine.9 ± 0. .2 24.3 ± 0.3 L. the as mean ± S.2 ± 0.8 ± 0.0 ± 0.0 mmol for potassium from 4.8 37.2 Grevillea robusta 21.3 14.0 mmol.5 ± 0.7 ± 0. potassium concentration in these nectars was 18.0 ± 0. tions for four species of Xylocopa flowers are only 1. The mean In an earlier survey of ion concentrations in nectar.4 ± 0. very rich nectar utilized by Xylocopa capitata is so low in cations that the error in calculating its energy content Results and Discussion would be only 0.3 11.0 ± 0.6 ± 0.4 ± 10.4 2. Ionic composition of nectars It is apparent from these results that honey bees have The results of the nectar survey of selected flowering a much wider choice of ion concentrations in nectar than plants are presented in Table 1.8 mmol for sodium and 4.3 mmol in Wachendorfia thyrsiflora (Haemodora.2 5. Sodium concentrations do carpenter bees.5 mmol in Callistemon capitata forages almost exclusively on Podalyria calyp- viminalis (Myrtaceae) to 21. with one excep- better (r = 0.3 Hiebert & Calder (1983) examined 19 species of flowers mmol.3 Fabaceae Indigofera langebergensis 0.4 5.5 ± 0.2 ± 0.6 ± 3.0 ± 0.2 Protea eximia 22.2 7.2 ± 0.7 ± 0.0 ± 0. In contrast.7 ± 4.3 ± 3.5 13. In the Cape Town area.8 Haemodoraceae Wachendorfia thyrsiflora 6.9 ± 7. Xylocopa of the nectar varied from 1. hummingbird flowers (Hiebert & Calder 1983: Table 1) ted anions) accounted for 4.8% of the apparent averaged 24. all different from the plant between sodium and sugar concentrations was somewhat families examined in the present work. Dierk. viminalis 9.4 9.5 ± 0.3 ± 0.2 Mimetes cucullatus 22.2 Myrtaceae Eucalyptus melliodora 5.6 ± 0. averaging 3.0 ± 0.3 ± 0.2 7.3 ± 0. cuneiforme 94.8 ± other members of the Fabaceae: nectar ion concentra- 1. sodium concentrations in the hummingbird robusta.3 12.0 ± 1.5 ± 0.3 59.94 S.2 7.7 ± 0. Roberts.l) (g/100 ml) (mmol) (mmol) Boraginaceae Echium fastuosum 2. ceae) to 68.6 ± 0. This would result in considerable errors in flowers.0 ± 0.2 ± 1.5 ± 0.4 mmol for the 18 species examined.2 7.58).0 ± 0. ficifolia 12.0 ± 0.4 5.1 27. (Nicolson 1990).5 ± 0. All results are presented Meyers.5 20.3 12.2 ± 0.2 ± 0.4 12.1 4.4 ± 0.2 14. while the correlation belonged to nine families. 1990. catherinae 19. Tsao & Wang 1980).9 ± 2. tion: Aloe (Liliaceae).2 1. ranging 0.3 4.4 8.13).5 ± 0.2 Chasmanthe floribunda 41. p < 0.1 41.4 8.1 68. with the highest cation contributions different from the values for bee flowers in Table 1.5 ± 0.9 ± centrations also showed considerable variation.5 1.4 ± 1.0 ± 1.5 ± 0.3 21. Potassium concentrations in these Sodium and potassium ions together (without associa.7 ± 0. being 14% in Callistemon viminalis and 10% in Grevillea However.0 ± 0.2 Proteaceae Leucospermum oleifolium 63.2 ± 0.7 ± 5.3 57.2 11.2 21.4 Iridaceae Watsonia meriana 11.4 ± 1.3 Callistemon citrinus 14. erubescens 76.3% (Nicolson 1990).3 E.1 ± 0.0 ± 0.2 13.1 39.4 16. These flowers poorly correlated (r = 0.-Afr.3 ± 0.0 mmol in two species of trata and two species of Virgilia. Na and K concentrations n = 10 for sugar concentrations.7 ± 2. Potassium and sugar concentrations were very known to be visited by hummingbirds.6 ± 1. Table 1 Nectar volume and composition of selected forage plants of honeybees Volume Sugar Na K Species (~J.3 55.4 L.

5 ± 0. Body fluid composition in relation to nectar ions however. with the limited data Leucospermum oleifolium.0 ± 0.3 39. Carpenter & Ziehl (1972). between bee. Leucospermum Na 21. Unusual- years.4 14. Nicolson 1990).7 ± 0.5 cited by Dadd 1968). However. Artificial tures and relative humidities were not measured either in food sources have been used extensively in research on the present study or in that of Hiebert & Calder (1983). and the present study) gives mean values of 5. Rebelo. Ambient tempera.8 mmol for sodium and 19. Species Ion Nectar Haemolymph Urine Nectar is generally derived from phloem sap (Durkee Echium Na 5. Similarly.7 ± 0. solutions to honey bees foraging at a feeder. for onion flowers have been omitted).5 over sodium in floral nectars is not unexpected. et a/.S.5 11. 25(2) 95 The only other information on nectar ions comes from sodium and potassium concentrations available to polli- an earlier study by Waller.7 mmol for potassium syndromes: 80% are pollinated by insects.5 1983). both the present nectar and the urine (Table 2). whereas honey Ranunculaceae.3 26. Levels of sodium in phloem sap are two orders of fastuosum K 11. Car- have been influenced by the plant families chosen for penter bees are solitary and forage only to satisfy their analysis (e.4 24. so the predominance of potassium melliodora K 16.4 37.g. not to provide a definitive analysis of the nectars. but it is clear that potassium concentrations are far higher than those of sodium.4 ± 6. and regres- Table 2 Ionic composition of body fluids of honey sion analysis shows that the concentrations of these two bees foraging on three different nectars ions in floral nectars are in fact slightly negatively correlated (r = 0. Combi.2 ± 0. nators.4 ± 5.7 ± 4. The values shown are slightly lower than the 47 mmol sodium and 27 mmol potassium given by Bee and bird pollination Florkin & Jeuniaux (1974) for the haemolymph of Apis Although there are differences in sugar concentration mellifica adults (subspecies not named). capitata. measured in the nectar of onion flowers by Waller et al. Zool. mean values sodium concentration (3-4 mmol. as described for dual bees to be less closely related in social species. They found that potassium Leucospermum oleifolium (Proteaceae) (Nicolson and content had a marked effect on the acceptability of sugar W. unpublished data). be compared directly with measurements made on floral nectars because honey production In honey bees foraging on three different plants with involves processing and concentrating of the nectar by nectars varying widely in ionic content. and in different seasons.3 18. Hiebert & Calder 1983.-Worswick. and potassium concentrations approximately three times Leucospermum.g.5 ± 1. who suggested that this reduced the attractive- measured at different times for Grevillea robusta and ness of onion flowers to bees. it is difficult to say. another species of Echium by Corbet (1978).6 magnitude lower than those of potassium (Ziegler 1962. trations differing greatly from those in Table 1 have been (1972). Eucalyptus Na 12.0 ± 0.and bird-pollinated flowers.6 ± in the south-western Cape according to their pollination 0. with one interesting exception.0 ± 0. Table 1 shows that sodium concentrations in some oleifolium K 8. which when foraging on extensively by honey bees when the latter are present at Virgilia divaricata forms a urine with the same low high densities (Rebelo 1987). Hiebert & Calder 1983).8 ± 0.2 ± 2. bees foraging study and that of Hiebert & Calder (1983) include on Eucalyptus melliodora produced urine with sodium flowers which are visited by both bees and birds. Pyke & In the case of bees foraging on Echium fastuosum and Waser 1981). Afr.9 ever.8 ± 0. Nicolson Siegfried & Oliver (1985) have classified the 426 species 1990. Hiebert & There is little evidence in the literature that honey Calder (1983) found differences in composition when bees may show preferences for different ion concentra- they compared nectars of some species in two successive tions in nectar. Firstly.0 ± 0. This contrasts with the Table 1. nectar bee foragers gather nectar for the colony as a whole.8 ± 0. There is great variation between species. J.7 ± 0. 1972.4 floral nectars are surprisingly high.4 ± 2. the feeding preferences of honey bees: one such experi- because in both cases the aim was to demonstrate the ment has shown that a colony need for water can affect . We sugar (and presumably ion) concentrations are greatly might expect nectar and urine concentrations of indivi- affected by environmental conditions.4 47.7 43. haemolymph sodium and potassium concentrations were closely regu- house bees.3 35. whether honey bee and hummingbird flowers ment between the levels of sodium and potassium in the differ in ionic characteristics. direct comparison of ionic or other constituents in the ning all the available data on nectar ion concentrations nectar of bee. there was reasonable agree- available. who analysed the honeystomach contents of honey bees The genus Erica would be excellent material for a foraging on a variety of crop and other plants. samples. Thirdly. Secondly. lated (Table 2). and 15% by (n = 51 species: the anomalous data obtained by Waller sunbirds.6 5. These data cannot. The tables on the mineral composition of honey Na and K concentrations in mmol presented by White (1975) show great variability among n = 6 for nectar. however. How. 10 for haemolymph and urine. (Waller et al. of which five species are included in as high as those in the nectar.5 ± 0. high potassium concentrations in the own needs and those of their offspring. 1990.0076). values for ly high levels of potassium (up to 300 mmol) were nectar volume and sugar.3 24.and bird-pollinated flowers (e. sodium and potassium concen. is a bird-pollinated genus which is utilized carpenter bee X.

) preliminary synthesis of pollination biology in the Cape Richards. Ecology dation for Research Development. 25(2) choice of sugar concentration (Ohguchi & Aoki 1983). and chloride in floral nectars: energy-free We are grateful for financial assistance from the Foun- contributions to refractive index and salt balance. In: measurement of evaporative water loss. Problems connected with inorganic Sci. Amer. 6: 139-170. 1983. 255-307. The effects of non-sugar nectar References constituents on estimates of nectar energy content. Syst. (eds) ideal to determine the ionic preferences of honey bees. Management implications.A. A. In: The biology of nectaries. exp. FAVRE. 1982. & CALDER.M. (ed. REBELO.) Crane. 1980. 1983. Simultaneous constitution and pollinator-plant coevolution. Ent.H. Vol. 157-206.. Ann. 222: 287-296. 1975. extrafloral nectaries. Y. J.W. BAKER. Sci. London. N. 193-211. 1985. & OLIVER. A. Amer. E. & LEDERHOUSE. 1983.M.. Press. 1981. 21-30. 1978. Academic Press. London. Princeton Bentley.G.. flora. PYKE. J.G. Sodium. E. M. Soclobiol. LEVINE. oxygen Coevolution of animals and plants.. Van Nostrand Reinhold. Rockstein. Effects of colony need BARROWS.. R. pp. J. 1974. In: The physiology of Insecta. N.. (ed. In: The pollination of flowers by insects.. Bull. 141. (ed. W. Sth.A.. pp. K. 1968. ent. New York & London. OHGUCHI. pp. Report No. New York. pp.M. Afr. Insect foraging energetics.G.E. J. & WANG.. Pollination syndromes of Erica species in the south- DURKEE.. & LOUW.S. 1987. 1975. CARPENTER. WALLER. Ecology 61: 992-996. and for permission 64: 399-402. potassium. carpenter bee.H.) on attractiveness of onion flowers to honey bees. C.C.J. Physiol. & Elias. Energetics of pollination. K. comprehensive survey.W. J. Studies of nectar NICOLSON.C. in press. Jones. S. 1974.. R. J. University of Texas Press. MEYERS. response to sodium chloride in females of a solitary bee. 1990. ARMS. Rev. (ed. Ecol. the carpenter bee Xylocopa capitata: water excess 372-374. A. S. In: A vulgare.A. 187-214. E. F. & AOKI. 97: 535-539..E. Composition of honey. pp. M. (eds) Gilbert.-Y. Hort. dilute nectars by hummingbird and honeyeater flowers.W. 5. New York. 100-140. E. Osmoregulation in a nectar-feeding swallowtail butterflies.D.T. S. & WASER. W. G. components of aqueous diets. H. 0. pp. S. B. FLORKIN. Honeybee ecology. 1985.96 S.. W. Prog. & consumption.G. G. Papilio glaucus.. T. BeluJv. In: Honey: a HEINRICH..-Afr. L.J.M. Bees and the nectar of Echium REBELO.B. A. and ion conservation. N. Acknowledgements HIEBERT. Dierk. Aggregation behavior and for water on optimal food choice in honey-bees. B. WHITE.. Nat. 14: 22-26. to sample bees and flowers in Kirstenbosch Botanic INOUYE.W. C. 1974.) Rebelo. Potassium in onion nectar and its probable effect composition. B. 1975. pp. 51: 270-280. 1983. G. D. Science 185: insect. 57: 189-193. O. P. SIEGFRIED. Biotropica 13: 260-270. L. In: Carefully designed experiments of this type would be Handbook of experimental pollination biology. T. 1990. Bot. .A.R. D.H.D. & Little. & JEUNIAUX. I. The production of Augochlora pura (Hymenoptera: Halictidae). Austin. HEINRICH. FEENY. ROBERTS. 12: 77--84.. Afr.H. R. Gardens. Soc.D. Academic Press. Ecol. P. Sodium: stimulus for puddling behaviour by tiger NICOLSON. Florida Ent. TSAO. and thoracic temperature during flight in a Raven. & BAKER. zool. (eds) SEELEY. 1-29. & ZIEHL. The ultrastructure of floral and western Cape. CORBET.N. LANUM. Hemolymph 1972. Columbia University University Press. S. DADD. Tydskr. Heinemann.G.