Honeybee Nests : Composition, Structure, Function

This work, a sequel to __Honeybees and Wax__ published nearly 30 years ago, starts with a brief introduction and discussion of nesting sites, their spaces and densities, self-organization of nest contents, and interspecific utilization of beeswax. The following chapters cover communication by vibrations and scents and wax secretion, and discuss the queen in relation to the combs. Discussions on completed nests include the significance of brood, the roles of pollen and nectar flow, and comb-building, and are followed by a triad of related chapters on the construction of cells and combs and their energetic costs. An in-depth examination of the conversion of wax scales into combs, the material properties of scale and comb waxes, and the wax gland complex are presented. The next chapters are devoted to a comprehensive analysis of the literature on the chemistry and synthesis of beeswax, and, finally, the material properties of honeybee silk are highlighted. Preface 6 Contents 10 1 General Introduction 16 Abstract 16 1.1...The Origin of Beeswax 17 1.2...Nests and Nesting 17 1.3...Self-Organization of Nest Contents 18 1.4...Interspecific Utilisation of Beeswax 18 1.5...Communication of Vibrations and Scents 19 1.6...Wax Secretion, Comb Construction and the Queen 19 1.7...The Significance of Brood 20 1.8...The Role of Pollen in Comb Construction 21 1.9...Nectar Flows and Comb-Building 21 1.10...Construction of Combs 22 1.11...Energetics of Honey/Beeswax Conversion 23 1.12...Construction of Cells 24 1.13...Conversion of Scale Wax into Combs 25 1.14...Material Properties of Scale and Comb Wax 25 1.15...The Wax Gland Complex 26 1.16...The Chemistry of Beeswax 27 1.17...Synthesis of Beeswax 27 1.18...Material Properties of Honeybee Silk 28 References 29 2 Nesting: Sites, Space and Density in Comb-Building 32 Abstract 32 2.1...Introduction 32 2.2...Nesting Sites 33 2.2.1 The Dwarf Honeybees 33 2.2.2 The Cavity-Nesting Honeybees 34 2.2.3 The Giant Honeybees 35 2.3...Nest Cavities 37 2.4...Colony Space and Density 39 2.4.1 Arrangement of Space 39 2.4.2 Density Versus Space 41 2.4.3 Reduction of Nest Size 43 2.4.4 Other Qualities of ‘‘Space’’ 45 2.5...Seasonality, Space and Density 47 References 49 3 Self-Organization of Nest Contents 55 Abstract 55 3.1...Introduction 55 3.2...Pattern Formation in Combs 57 3.2.1 Reaction--Diffusion Systems Pattern Formation 57 3.2.2 Template Effects? 60 3.2.3 Recent Models 61 3.3...Developmental Cycles of Apis florea Nests 64 References 68 4 Intraspecific and Interspecific Comb-Building 70 Abstract 70 4.1...Introduction 70 4.2...Intraspecific Comb Wax Salvage 71 4.3...Interspecific Wax Salvage 73 4.4...Interspecific Wax Discrimination 75 4.5...Comb-Building in Mixed-Species Colonies 79 4.5.1 Organisation of Mixed-Species Colonies and Wax Foundation 79 4.5.2 Cell-Size and Wax Discrimination 80 4.5.3 Cell-Size Modification of Foundation Sheets 80 4.5.4 Freely-Built Combs 82 4.5.5 Utilisation of the Newly Built Combs 86 4.5.6 General Comb-Building 86 4.5.7 Comb-Building in Mixed-Species Colonies 87 References 88 5 Communication by Vibrations and Scents in the Comb 92 Abstract 92 5.1...Introduction 92 5.2...Vibrations 94 5.2.1 Queen Honeybees 95 5.2.2 Worker Vibrations 98 5.3...Scents 106 5.3.1 Waggle Dance Scent-marking: Probable Cause? 107 5.3.2 Comb and Scents 108 5.3.3 Capping Brood Cells 110 References 114 6 Wax Secretion, Comb Construction and the Queen 117 Abstract 117 6.1...Introduction 117 6.2...The Queen: A Necessary Stimulus for Comb-Building? 119 6.3...Comb-Building Experiments by Whiffler and Hepburn (1991a) 120 6.3.1 Queenright and Queenless Colonies 120 6.3.2 Free-Running and Confined Queens 122 6.3.3 Division Board Experiments 122 6.3.4 General Conclusions from the Experiments of Whiffler and Hepburn (1991a, b) 126 6.4...Comb-Building Experiments of Ledoux et al. (2001) 128 6.5...Perception of Queenrightness 129 6.6...Comb-Building Experiments of Maisonnasse et al. (2010) 134 6.7...The Construction of Queen Cells 135 References 139 7 The Significance of Brood 143 Abstract 143 7.1...Introduction 143 7.2...Parthenogenesis 144 7.3...Oviposition by Queens 145 7.4...The Meaning of Brood 146 7.5...Efficacy of Open Brood 149 7.6...Drone Brood 152 7.7...Brood-Rearing and Honey Storage 153 References 154 8 The Role of Pollen in Honeybee Colonies 156 Abstract 156 8.1...Pollen and Brood 156 8.2...Pollen and Wax Production 160 8.3...Physical Presence and Regulation of Pollen in the Colony 164 8.4...Pollen Pheromones 168 8.5...Pattern and Function of Pollen Cells 171 8.6...Cell Allocation 173 References 181 9 Nectar Flows and Comb-Building 185 Abstract 185 9.1...Introduction 185 9.2...Temperate Zone Spring as a Stimulus 187 9.3...Tropical Areas: Environmental-Based Construction 189 9.4...Nectar, the Unqualified Stimulus for Comb Construction 193 9.4.1 Hoarding Assays 195 9.4.2 The Honey Stomach 199 9.5...Decision-Making and Regulation of Comb-Building 199 9.6...Who are the Comb Builders? 202 9.7...Nectar Intake and Comb Fullness 207 9.8...Termination of the Stimulus 209 References 213 10 Construction of Combs 217 Abstract 217 10.1...Introduction 217 10.2...Parallelism Between Combs 218 10.3...Festoons and Torsion 219 10.4...Festoons and Comb Growth 223 10.5...Evidence of a Sense of Equilibrium 225 10.6...Application of the Sense of Equilibrium 227 10.7...The Orientation of Combs 229 10.8...Behavioural Aspects of Comb Construction 230 References 230 11 Energetics of Honey/Beeswax Conversion 232 Abstract 232 11.1...Introduction 232 11.2...Cumulative Ratios 234 11.3...Measures of Conversion Efficiency 239 11.4...Temperature and Wax Production 244 References 244 12 Construction of Cells 246 Abstract 246 12.1...Introduction 246 12.2...Manipulation of Wax Scales 247 12.3...Comb Operations 247 12.4...Inception of the Nest 250 12.5...Recognition of Cell Patterns 257 12.6...Assessment of Cell Size 259 12.7...The Cell Base: Changing from Rhombus to Hemisphere 263 References 265 13 Conversion of Wax Scales into Comb Wax 268 Abstract 268 13.1...Introduction 268 13.2...Wax Scales 270 13.3...Chemical Differences Between Scale and Comb Wax 270 13.4...Maturation of Newly Constructed Combs 271 13.5...Wax Scales 275 13.6...Unnatural Building Materials 276 14 Material Properties of Scale and Comb Wax 282 Abstract 282 14.1...Introduction 282 14.2...Temperature Effects 283 14.3...Crystal Changes 286 14.4...Tensile Properties 289 14.5...Crystal Texture 294 14.6...Wax Proteins 294 14.7...aacgr-Helical Silk 296 14.8...Optical Studies 297 14.9...X-ray Diffraction Studies 298 14.10...Crystallites of Beeswax 300 14.11...Origins of Crystallites in Beeswax 304 References 305 15 The Wax Gland Complex 309 Abstract 309 15.1...Introduction 309 15.2...Source of Secretion 313 15.2.1 The Cuticle 313 15.2.2 The Epidermis 315 15.2.3 Fat Body and Oenocytes 317 15.2.4 Synchronising Cellular Activity 319 15.2.5 Ultrastructure of the Organelles of Wax Gland Cells 321 References 325 16 The Chemistry of Beeswax 327 Abstract 327 16.1...Introduction 327 16.2...Chemical Composition 328 16.3...Chemometrics 331 16.3.1 Chemometric Classification of Beeswaxes 333 16.3.2 Discrimination and Classification of Beeswaxes 338 16.4...The Proteins of Beeswax 342 16.5...Plant-Derived Aromatic Volatiles and Colourants in Beeswax 343 References 345 17 Synthesis of Beeswax 348 Abstract 348 17.1...Introduction: Proof of Beeswax Synthesis 348 17.1.1 François Huber (1814) 349 17.1.2 The Chemists: Dumas and Edwards (1843) 350 17.2...Routes of Synthesis 351 17.3...Biochemical Investigations on Beeswax Synthesis 351 17.3.1 Hypothetical Scheme for Beeswax Synthesis 351 17.3.2 Monoester Synthesis 354 17.3.3 Cuticular and Comb Waxes 355 17.4...Cellular Basis of Synthesis 358 17.4.1 Chemical Composition and the Ages of Worker Bees 360 17.5...Secretion of Beeswax 362 References 369 18 Material Properties of Honeybee Silk 373 Abstract 373 18.1...Introduction 373 18.2...Honeybee Silk: An aacgr-Helical Protein 375 18.3...Behaviour of Silk at Different Temperatures 377 18.4...Relative Crystallinity 378 18.5...Solvent Effects on Silk 379 18.6...Honeybee Silk: An aacgr-Helical Silk and a Coiled-Coil Protein 381 18.7...Molecular Dynamics of aacgr-Helical Proteins 383 18.8...Genetic Basis of Honeybee aacgr-Helical Fibroin 385 Index 389 "This work, a sequel to Honeybees and Wax published nearly 30 years ago, starts with a brief introduction and discussion of nesting sites, their spaces and densities, self-organization of nest contents, and interspecific utilization of beeswax The following chapters cover communication by vibrations and scents and wax secretion, and discuss the queen in relation to the combs Discussions on completed nests include the significance of brood, the roles of pollen and nectar flow, and comb-building, and are followed by a triad of related chapters on the construction of cells and combs and their energetic costs An in-depth examination of the conversion of wax scales into combs, the material properties of scale and comb waxes, and the wax gland complex are presented The next chapters are devoted to a comprehensive analysis of the literature on the chemistry and synthesis of beeswax, and, finally, the material properties of honeybee silk are highlighted - Font no determinada Front Matter....Pages i-xvi General Introduction....Pages 1-16 Nesting: Sites, Space and Density in Comb-Building....Pages 17-39 Self-Organization of Nest Contents....Pages 41-55 Intraspecific and Interspecific Comb-Building....Pages 57-78 Communication by Vibrations and Scents in the Comb....Pages 79-103 Wax Secretion, Comb Construction and the Queen....Pages 105-130 The Significance of Brood....Pages 131-143 The Role of Pollen in Honeybee Colonies....Pages 145-173 Nectar Flows and Comb-Building....Pages 175-206 Construction of Combs....Pages 207-221 Energetics of Honey/Beeswax Conversion....Pages 223-236 Construction of Cells....Pages 237-258 Conversion of Wax Scales into Comb Wax....Pages 259-272 Material Properties of Scale and Comb Wax....Pages 273-299 The Wax Gland Complex....Pages 301-318 The Chemistry of Beeswax....Pages 319-339 Synthesis of Beeswax....Pages 341-365 Material Properties of Honeybee Silk....Pages 367-382 Back Matter....Pages 383-389 General Introduction -- Nesting : Sites, Space And Density In Comb-building -- Self-organization Of Nest Contents -- Intraspecific And Interspecific Comb-building -- Communication By Vibrations And Scents In The Comb -- Wax Secretion, Comb Construction And The Queen -- The Significance Of Brood -- The Role Of Pollen In Honeybee Colonies -- Nectar Flows And Comb-building -- Construction Of Combs -- Energetics Of Honey/beeswax Conversion -- Construction Of Cells -- Conversion Of Wax Scales Into Comb Wax -- Material Porperties Of Scale And Comb Wax -- The Wax Gland Complex -- The Chemistry Of Beeswax -- Synthesis Of Beeswax -- Material Properties Of Honeybee Silk. H.r. Hepburn, C.w.w. Pirk, O. Duangphakdee. Revised And Updated Sequel To: Honeybees And Wax : An Experimental Natural History / H.r. Hepburn. ©1986. (preface And Page [4] Of Cover) Includes Bibliographical References And Index.