Obituary Professor Sue Nicolson 1950-2023: Sweet solutions: pollinators and their physiology




On 27 April 2023 the scientific community lost an inspirational and influential biologist of rare quality following the death of Professor Sue Nicolson.


Altaye SZ, Pirk CWW, Crewe RM, Nicolson SW (2010) Convergence of carbohydrate-biased intake targets in caged worker honeybees fed different protein sources. Journal of Experimental Biology 213:3311-3318. DOI:

Archer CR, Koehler A, Pirk CWW, Oosthuizen V, Apostolides Z, Nicolson SW (2014a) Antioxidant supplementation can reduce the survival costs of excess amino acid intake in honeybees. Journal of Insect Physiology 71:78-86. DOI:

Archer CR, Pirk CWW, Carvalheiro LG, Nicolson SW (2014b) Economic and ecological implications of geographic bias in pollinator ecology in the light of pollinator declines. Oikos 123:401-407. DOI:

Archer CR, Pirk CWW, Wright GA, Nicolson SW (2014c) Nutrition affects survival in African honeybees exposed to interacting stressors. Functional Ecology 28:913-923. DOI:

Barnes K, Nicolson SW, Van Wyk BE (1995) Nectar sugar composition in Erica. Biochemical Systematics and Ecology 23:419-423. DOI:

Carvalheiro LG, Seymour CL, Nicolson SW, Veldtman R (2012) Creating patches of native flowers facilitates crop pollination in large agricultural fields: mango as a case study. Journal of Applied Ecology 49:1373-1383. DOI:

Carvalheiro LG, Seymour CL, Veldtman R, Nicolson SW (2010) Pollination services decline with distance from natural habitat even in biodiversity-rich areas. Journal of Applied Ecology 47:810-820. DOI:

Carvalheiro LG, Veldtman R, Shenkute AG, Tesfay GB, Pirk CWW, Donaldson JS, Nicolson SW (2011) Natural and within-farmland biodiversity enhances crop productivity. Ecology Letters 14:251-259. DOI:

Chown SL, Nicolson S (2004) Insect Physiological Ecology: Mechanisms and Patterns. Oxford University Press, Oxford, UK DOI:

Demares FJ, Crous KL, Pirk CWW, Nicolson SW, Human H (2016) Sucrose sensitivity of honey bees Is differently affected by dietary protein and a neonicotinoid pesticide. Plos One 11. DOI:

Demares FJ, Pirk CWW, Nicolson SW, Human H (2018) Neonicotinoids decrease sucrose responsiveness of honey bees at first contact. Journal of Insect Physiology 108:25-30. DOI:

Demares FJ, Yusuf AA, Nicolson S, Pirk CWW (2017) Effect of brood pheromone on survival and nutrient intake of African honey bees (Apis mellifera scutellata) under controlled conditions. Journal of Chemical Ecology 43:443-450. DOI:

du Rand EE, Pirk CWW, Nicolson SW, Apostolides Z (2017) The metabolic fate of nectar nicotine in worker honey bees. Journal of Insect Physiology 98:14-22. DOI:

du Rand EE, Smit S, Beukes M, Apostolides Z, Pirk CWW, Nicolson SW (2015) Detoxification mechanisms of honey bees (Apis mellifera) resulting in tolerance of dietary nicotine. Scientific Reports 5. DOI:

du Rand EE, Stutzer C, Human H, Pirk CWW, Nicolson SW (2020) Antibiotic treatment impairs protein digestion in the honeybee, Apis mellifera. Apidologie 51:94-106. DOI:

Ellis MB, Nicolson SW, Crewe RM, Dietemann V (2008) Hygropreference and brood care in the honeybee (Apis mellifera). Journal of Insect Physiology 54:1516-1521. DOI:

Ellis MB, Nicolson SW, Crewe RM, Dietemann V (2010) Brood comb as a humidity buffer in honeybee nests. Naturwissenschaften 97:429-433. DOI:

Fielding PJ, Nicolson SW (1980) Regulation of haemolymph osmolarity and ions in the green protea beetle, Trichostetha fascicularis, during dehydration and rehydration. Comparative Biochemistry and Physiology 67A:691-694. DOI:

Fleming PA, Hofmeyr SD, Nicolson SW, du Toit JT (2006) Are giraffes pollinators or flower predators of Acacia nigrescens in Kruger National Park, South Africa? Journal of Tropical Ecology 22:247-253. DOI:

Fleming PA, Nicolson SW (2002) How important is the relationship between Protea humiflora (Proteaceae) and its non-flying mammal pollinators? Oecologia 132:361-368. DOI:

Fleming PA, Xie S, Napier K, McWhorter TJ, Nicolson SW (2008) Nectar concentration affects sugar preferences in two Australian honeyeaters and a lorikeet. Functional Ecology 22:599-605. DOI:

Franke E, Jackson S, Nicolson S (1998) Nectar sugar preferences and absorption in a generalist African frugivore, the Cape White-eye Zosterops pallidus. Ibis 140:501-506. DOI:

Hanrahan SA, Nicolson SW (1987) Ultrastructure of the Malpighian tubules of Onymacris plana plana Peringuey (Coleoptera: Tenebrionidae). International Journal of Insect Morphology & Embryology 16:99-119. DOI:

Human H, Archer CR, du Rand EE, Pirk CWW, Nicolson SW (2014) Resistance of developing honeybee larvae during chronic exposure to dietary nicotine. Journal of Insect Physiology 69:74-79. DOI:

Human H, Nicolson SW (2006) Nutritional content of fresh, bee-collected and stored pollen of Aloe greatheadii var. davyana (Asphodelaceae). Phytochemistry 67:1486-1492. DOI:

Human H, Nicolson SW, Dietemann V (2006) Do honeybees, Apis mellifera scutellata, regulate humidity in their nest? Naturwissenschaften 93:397-401. DOI:

Human H, Nicolson SW, Strauss K, Pirk CWW, Dietemann V (2007) Influence of pollen quality on ovarian development in honeybee workers (Apis mellifera scutellata). Journal of Insect Physiology 53:649-655. DOI:

Jackson S, Nicolson SW (2002) Xylose as a nectar sugar: from biochemistry to ecology. Comparative Biochemistry and Physiology B-Biochemistry & Molecular Biology 131:613-620. DOI:

Jackson S, Nicolson SW, Lotz CN (1998a) Sugar preferences and "side bias" in Cape Sugarbirds and Lesser Double-collared Sunbirds. Auk 115:156-165. DOI:

Jackson S, Nicolson SW, van Wyk BE (1998b) Apparent absorption efficiencies of nectar sugars in the Cape Sugarbird, with a comparison of methods. Physiological Zoology 71:106-115. DOI:

Johnson SA, Jackson S, Abratt VR, Wolfaardt GM, Cordero-Otero R, Nicolson SW (2006a) Xylose utilization and short-chain fatty acid production by selected components of the intestinal microflora of a rodent pollinator (Aethomys namaquensis). Journal of Comparative Physiology B-Biochemical Systems and Environmental Physiology 176:631-641. DOI:

Johnson SA, Nicolson SW, Jackson S (2006b) Nectar xylose metabolism in a rodent pollinator (Aethomys namaquensis): Defining the role of gastrointestinal microflora using 14C-labeled xylose. Physiological and Biochemical Zoology 79:159-168. DOI:

Johnson SA, van Tets IG, Nicolson SW (1999) Sugar preferences and xylose metabolism of a mammal pollinator, the Namaqua rock mouse (Aethomys namaquensis). Physiological and Biochemical Zoology 72:438-444. DOI:

Köhler A, Leseigneur CDC, Verburgt L, Nicolson SW (2010) Dilute bird nectars: viscosity constrains food intake by licking in a sunbird. American Journal of Physiology-Regulatory Integrative and Comparative Physiology 299:R1068-R1074. DOI:

Köhler A, Pirk CWW, Nicolson SW (2012a) Honeybees and nectar nicotine: Deterrence and reduced survival versus potential health benefits. Journal of Insect Physiology 58:286-292. DOI:

Köhler A, Pirk CWW, Nicolson SW (2012b) Simultaneous stressors: Interactive effects of an immune challenge and dietary toxin can be detrimental to honeybees. Journal of Insect Physiology 58:918-923. DOI:

Köhler A, Raubenheimer D, Nicolson SW (2012c) Regulation of nutrient intake in nectar-feeding birds: insights from the geometric framework. Journal of Comparative Physiology B-Biochemical Systems and Environmental Physiology 182:603-611. DOI:

Köhler A, Verburgt L, Fleming PA, Nicolson SW (2008) Changes in nectar concentration: how quickly do whitebellied sunbirds (Cinnyris talatala) adjust feeding patterns and food intake? Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology 178:785-793. DOI:

Köhler A, Verburgt L, Nicolson SW (2006) Short-term energy regulation of whitebellied sunbirds (Nectarinia talatala): effects of food concentration on feeding frequency and duration. Journal of Experimental Biology 209:2880-2887. DOI:

Leon B, Nicolson SW (1997) Metabolic rate and body temperature of an African sunbird, Nectarinia chalybea: daily rhythm and the effect of ambient temperature. South African Journal of Zoology 32:31-36 DOI:

Lotz CN, Nicholson SW (1996) Sugar preferences of a nectarivorous passerine bird, the Lesser Double-collared Sunbird (Nectarinia chalybea). Functional Ecology 10:360-365. DOI:

Lotz CN, Nicolson SW (1999) Energy and water balance in the lesser double-collared sunbird (Nectarinia chalybea) feeding on different nectar concentrations. Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology 169:200-206. DOI:

Louw GN, Nicolson SW (1983) Thermal, energetic and nutritional considerations in the foraging and reproduction of the carpenter bee Xylocopa capitata. African Entomology (formely Journal of the Entomological Society of Southern Africa) 46:227-240

Louw GN, Nicolson SW, Seely MK (1986) Respiration beneath desert sand: carbon dioxide diffusion and respiratory patterns in a tenebrionid beetle. Journal of Experimental Biology 120:443-447 DOI:

Moloney CL, Nicolson SW (1984) Water relations and haemolymph composition of two intertidal spiders (Order Araneae). Journal of Experimental Marine Biology and Ecology 83:275-284. DOI:

Nepi M, Human H, Nicolson S, Cresti L, Pacini E (2006) Nectary structure and nectar presentation in Aloe castanea and A. greatheadii var. davyana (Asphodelaceae). Plant Systematics and Evolution 257:45-55. DOI:

Nicolson S, P.M. H, Gardiner BOC, Maddrell SH (1974) Effects of starvation and dehydration on osmotic and ionic balance in Carausius morosus. Journal of Insect Physiology 20:2061-2069. DOI:

Nicolson SW (1972) Osmoregulation in larvae of New Zealand salt-water mosquito Opifex fuscus Hutton Journal of Entomology Series (A) - General Entomology 47:101-108 DOI:

Nicolson SW (1976a) Diuresis in the cabbage white butterfly, Pieris brassicae: fluid secretion by the Malpighian tubules. Journal of Insect Physiology 22:1347-1356. DOI:

Nicolson SW (1976b) Diuresis in the cabbage white butterfly, Pieris brassicae: water and ion regulation and the role of the hindgu. Journal of Insect Physiology 22:1623-1630. DOI:

Nicolson SW (1976c) The hormonal control of diuresis in the cabbage white butterfly Pieris brassicae. Journal of Experimental Biology 65:565-575 DOI:

Nicolson SW (1980) Water balance and osmoregulation in Onymacris plana, a tenebrionid beetle from the Namib Desert. Journal of Insect Physiology 26:315-320. DOI:

Nicolson SW (1987) Absence of endothermy in flightless dung beetles from southern Africa. South African Journal of Zoology 22:323-324 DOI:

Nicolson SW (1990) Osmoregulation in a nectar-feeding insect, the carpenter bee Xylocopa capitata: water excess and ion conservation. Physiological Entomology 15:433-440 DOI:

Nicolson SW (1991) Diuresis or clearance: is there a physiological role for the "diuretic hormone" of the desert beetle Onymacris? Journal of Insect Physiology 37:447-452. DOI:

Nicolson SW (1993a) The ionic basis of fluid secretion in insect Malpighian tubules: advances in the last ten years. Journal of Insect Physiology 39:451-458. DOI:

Nicolson SW (1993b) Low nectar concentrations in a dry atmosphere: a study of Grevillea robusta (Proteaceae) and Callistemon viminalis (Myrtaceae). South African Journal of Science 89:473-477

Nicolson SW (1994a) Eucalyptus nectar: production, availability, composition and osmotic consequences for the larva of the eucalypt nectar fly, Drosophila flavohirta. South African Journal of Science 90:75-79

Nicolson SW (1994b) Pollen feeding in the eucalypt nectar fly, Drosophila flavohirta. Physiological Entomology 19:58-60. DOI:

Nicolson SW (1995) Direct demonstration of nectar reabsorption in the flowers of Grevillea robusta (Proteaceae). Functional Ecology 9:584-588. DOI:

Nicolson SW (2006) Water management in nectar-feeding birds. American Journal of Physiology-Regulatory Integrative and Comparative Physiology 291:R828-R829. DOI:

Nicolson SW (2009) Water homeostasis in bees, with the emphasis on sociality. Journal of Experimental Biology 212:429-434. DOI:

Nicolson SW (2011) Bee food: the chemistry and nutritional value of nectar, pollen and mixtures of the two. African Zoology 46:197-204. DOI:

Nicolson SW (2022) Sweet solutions: nectar chemistry and quality. Philosophical Transactions of the Royal Society B-Biological Sciences 377. DOI:

Nicolson SW, Bartholomew GA, Seely MK (1984a) Ecological correlates of locomotion speed, morphometrics and body temperature in three Namib Desert tenebrionid beetles. South African Journal of Zoology 19:131-134 DOI:

Nicolson SW, de Veer L, Koehler A, Pirk CWW (2013) Honeybees prefer warmer nectar and less viscous nectar, regardless of sugar concentration. Proceedings of the Royal Society B-Biological Sciences 280. DOI:

Nicolson SW, Hanrahan SA (1986) Diuresis in a desert beetle? Hormonal control of the Malpighian tubules of Onymacris plana (Coleoptera: Tenebrionidae). Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology 156:407-413. DOI:

Nicolson SW, Human H, Pirk CWW (2022) Honey bees save energy in honey processing by dehydrating nectar before returning to the nest. Scientific Reports 12. DOI:

Nicolson SW, Leader JP (1974) The permeability to water of the cuticle of the larva of Opifex fuscus (Hutton) (Diptera, Culicidae). Journal of Experimental Biology 60:593-603 DOI:

Nicolson SW, Louw GN (1980) Preflight thermogenesis, conductance and thermoregulation in the protea beetle, Trichostetha fascicularis (Scarabaeidae: Cetoniinae). South African Journal of Science 76:124-126

Nicolson SW, Louw GN (1982) Simultaneous measurement of evaporative water loss, oxygen consumption, and thoracic temperature during flight in a carpenter bee. Journal of Experimental Zoology 222:287-296. DOI:

Nicolson SW, Louw GN, Edney EB (1984b) Use of a ventilated capsule and tritiated water to measure evaporative water losses in a tenebrionid beetle. Journal of Experimental Biology 108:477-481 DOI:

Nicolson SW, Nepi M (2005) Dilute nectar in dry atmospheres: Nectar secretion patterns in Aloe castanea (Asphodelaceae). International Journal of Plant Sciences 166:227-233. DOI:

Nicolson SW, Nepi M, Panini E (2007) Nectaries and nectar. Springer, Dordrecht. DOI:

Nicolson SW, Van Wyk BE (1998) Nectar sugars in Proteaceae: Patterns and processes. Australian Journal of Botany 46:489-504. DOI:

Nicolson SW, Worswick PVW (1990) Sodium and potassium concentrations in floral nectars in relation to foraging by honey bees. South African Journal of Zoology 25:93-96 DOI:

Nicolson SW, Wright GA (2017) Plant-pollinator interactions and threats to pollination: perspectives from the flower to the landscape. Functional Ecology 31:22-25. DOI:

Paoli PP, Donley D, Stabler D, Saseendranath A, Nicolson SW, Simpson SJ, Wright GA (2014) Nutritional balance of essential amino acids and carbohydrates of the adult worker honeybee depends on age. Amino Acids 46:1449-1458. DOI:

Pirk CWW, Boodhoo C, Human H, Nicolson S (2010) The importance of protein type and protein to carbohydrate ratio for survival and ovarian activation of caged honeybees (Apis mellifera scutellata). Apidologie 41:62-72. DOI:

Robertson HG, Nicolson SW, Louw GN (1982) Osmoregulation and temperature effects on water loss and oxygen consumption in two species of African scorpion. Comparative Biochemistry and Physiology a-Physiology 71:605-609. DOI:

Shi L, Nicolson SW, Yang Y, Wu J, Yan S, Wu Z (2020) Drinking made easier: honey bee tongues dip faster into warmer and/or less viscous artificial nectar. Journal of Experimental Biology 223. DOI:

Stabler D, Paoli PP, Nicolson SW, Wright GA (2015) Nutrient balancing of the adult worker bumblebee (Bombus terrestris) depends on the dietary source of essential amino acids. Journal of Experimental Biology 218:793-+. DOI:

Stevenson PC, Koch H, Nicolson SW, Brown MJF (2022) Natural processes influencing pollinator health. Philosophical Transactions of the Royal Society B-Biological Sciences 377. DOI:

Stevenson PC, Nicolson SW, Wright GA (2017) Plant secondary metabolites in nectar: impacts on pollinators and ecological functions. Functional Ecology 31:65-75. DOI:

van Tets IG, Hutchings K, Nicolson SW (2000) Relative merits of using purified proteins and natural foods in studies of wildlife nutrition: the nitrogen requirements of a rodent pollinator. African Zoology 35:207-215 DOI:

van Tets IG, Nicolson SW (2000) Pollen and the nitrogen requirements of the lesser double-collared sunbird. Auk 117:826-830.[0826:Patnro]2.0.Co;2 DOI:

Van Wyk BE, Nicolson SW (1995) Xylose is a major nectar sugar in Protea and Faurea. South African Journal of Science 91:151-153

Vanbergen AJ, Baude M, Biesmeijer JC, Britton NF, Brown MJF, Brown M, Bryden J, Budge GE, Bull JC, Carvell C, Challinor AJ, Connolly CN, Evans DJ, Feil EJ, Garratt MP, Greco MK, Heard MS, Jansen VAA, Keeling MJ, Kunis WE, Marris GC, Memmott J, Murray JT, Nicolson SW, Osborne JL, Paxton RJ, Pirk CWW, Polce C, Potts SG, Priest NK, Raine NE, Roberts S, Ryabov EV, Shafir S, Shirley MDF, Simpson SJ, Stevenson PC, Stone GN, Termansen M, Wright GA, Insect Pollinators I (2013) Threats to an ecosystem service: pressures on pollinators. Frontiers in Ecology and the Environment 11:251-259. DOI:

Wei J, Rico-Guevara A, Nicolson SW, Brau F, Damman P, Gorb SN, Wu Z, Wu J (2023) Honey bees switch mechanisms to drink deep nectar efficiently. Proceedings of the National Academy of Sciences 120:e2305436120. DOI:

Withers P, Louw G, Nicolson S (1982) Water loss, oxygen consumption and colour change in "waterproof" reed frogs (Hyperolius). South African Journal of Science 78:30-32

Wright G, Shafir S, Nicolson S, Stevenson P (2022) Bee nutrition. Apix Biosciences

Wright GA, Nicolson SW, Shafir S (2018) Nutritional Physiology and Ecology of Honey Bees. In: Berenbaum MR (ed) Annual Review of Entomology, Vol 63, vol 63, pp 327-344 DOI:




How to Cite

Pirk, C., Crewe, R., & Stevenson, P. (2024). Obituary Professor Sue Nicolson 1950-2023: Sweet solutions: pollinators and their physiology . Journal of Pollination Ecology, 35, 104–111.



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