honeybees and wax ||
TRANSCRIPT
H. R. Hepburn
Honeybees and Wax An Experimental Natural History
With 82 Figures
Springer-Verlag Berlin Heidelberg New York
London Paris Tokyo
Prof. Dr. H. R. HEPBURN
Department of Physiology University of the Witwatersrand
York Road, Parktown 2193 Johannesburg
South Africa
Current Address: Department of Zoology and Entomology
Rhodes University Grahamstown 6140
South Africa
Cover illustration Comb with false cells built by the African honeybee, drawn by C. P. Richards
(originally published in South African Bee Journal 1983)
ISBN-13: 978-3-642-71460-3 001: 10.1007/978-3-642-71458-0
e-ISBN-13: 978-3-642-71458-0
Library of Congress Cataloging-in-Publication Data. Hepburn, H. R. Honeybees and wax. Bibliography: p. Includes index. 1. Honeybee. 2. Bees wax. I. Title. QL568.A6H395
1986 638'.17 86-17899
This work is subject to copyright. All rights are reserved, whether th« whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under §54 of the German Copyright Law where copies are made for other
than private use a fee is payable to "Verwertungsgesellschaft Wort", Munich.
© Springer-Verlag Berlin Heidelberg 1986 Softcover reprint of the hard cover I st edition 1986
The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
213113130-543210
For Professor M. S. Blum
PREFACE
"Instead of dirt and poison we have rather chosen to fill our hives with honey
and wax; thus furnisning mankind with the two noblest of things, which are
sweetness and light". Mindful of Swift's dictum, this compilation is offered as an
exhaustive coverage of a smallish literature on the synthesis and secretion of
beeswax, its elaboration into combs and the factors which bear on the execution
of these processes by honeybees.
To codify any aspect of the biology of an animal of agricultural importance is to
sift through myriad observations and experiments, centuries old, that come down
to us enshrouded in the folk literature. It is evident that wars and languages
have also acted as barriers to the dissemination of knowledge about honeybees.
Thus, particular care has been given to the primacy of discovery and its con
textual significance.
I have endeavoured to not over-interpret data and to allow the authors' works to
speak for themselves. I have also tried to indicate some of the more obvious gaps
in our knowledge of honeybees in relation to wax and to suggest some directions
as to where we might proceed, aided by discoveries made on other animals and
plants. This was done to remind the seasoned bee-hand of our general neglect
of beeswax biology, historically constituting less than a percentage point of the
apicultural literature. And, hopefully, to present a comprehensive literature
survey to young scientists in other fields who might be casting around for the
magic combination of the accessible problem and the suitable experimental animal.
The reader is reminded that the wax of bees and the bees and their wax are rather
specific aspects of the life of the honeybee colony. Vast amounts of more general
information on honeybee biology can be found in the fine works of Ribbands (1953), BUdel and Herold (1960), Chauvin (1968), Butler (1974), Dadant and Sons
(1975) and Free (1977). Similarly, there is a smaller technical and practical lit
erature on the beeswax of commerce that includes the works of Otto (1944), Root
(1951), Warth (1956), Bull (1959-1970) and Coggshall and Morse (1984).
It is a pleasure to record my indebtedness to colleagues who have assisted in the
production of this book. R. Darchen, M.F. Johannsmeier, R.A. Morse, S.W.
Nicholson, G. Pretorius, T. Robinson, S. Taber and A. P. Tulloch read or dis-
VIII
cussed parts of the manuscript and made suggestions for its improvement.
Needless to say, the faults remaining are solely mine. M. DrUsedau assisted with
the translations from German. M.J.M. Nijland in particular and P.A. Middleton
and E.J. Hepburn processed the many revisions of the text. C. P. Richards has
again kindly provided all of the necessary drawings. The librarians of the
International Bee Research Association, the State Library at Pretoria and the
University of the Witwatersrand were persistent and successful in procuring the
necessary literature. M. Locke kindly provided some photomicrographs. Other
sources of borrowed material are indicated in figure legends and tables. Both the
Council for Scientific and Industrial Research and the Plant Protection Research
I nstitute have generously supported my own research on honeybees.
Finally, I wish to record that this book is a consequence of my having been
stimulated by the works of C.G. Butler, J.B. Free and C.R. Ribbands.
H.R. Hepburn
CHAPTER 1
Introduction
CONTENTS
PART I: THE NATURE AND PRODUCTION OF BEESWAX
CHAPTER 2
The Origin of Beeswax: an Historical Perspective
The Classical Era
The Renaissance
The Enlightenment
CHAPTER 3
Sou rce of Secretion
The Cuticle: the Search for Holes
Filaments or Microtubules?
Tracheation
The Epidermis
Fat Body and Oenocytes
Synchronising Cellular Activity
CHAPTER 4
Rise and Fall of the Epithelium
Age and the Division of Labour
Rejuvenation of Old Bees
Newly Settled Swarms
Laying Workers
Racial Differences
Tragedies and Disease
Winter
Cell Height and Secretory Performance
CHAPTER 5
Composition and Synthesis of Beeswax
. 5
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Gross Composition of Beeswax
The Proteins of Beeswax
Proof of Synthesis
Routes of Synthesis
Recent Refinements
CHAPTER 6
Food Conversion Ratios in the Production of Beeswax
The Agricultu ral Chemists
Cumulative Ratios
Direct Costs of Wax Production
Measures of Conversion Efficiency
Temperatu re and Wax Production
PART II: THE MANIPULATION OF WAX BY HONEYBEES
CHAPTER 7
Manipulation of Wax Scales
Removal of the Wax Scales (Casteel 1912)
The Scale-Removing Organ(Casteel 1912)
Mandibulation of the Scales
Free Scales (Casteel 1912)
Partial Removal of Scales(Casteel 1912)
Producers and Builders(Casteel 1912)
CHAPTER 8
Metamorphosis of Wax
The Structure of Virgin Scale Wax
Optical and Diffraction Studies
Crystallography
Mechanical Properties of Wax
Matu ration of Newly Constructed Combs
Foreign Substances
CHAPTER 9
The Construction of Cells
Minor Building Operations
I nception of the Nest
The Irregular Nature of Cells
Recognition of Cell Patterns
Interlude: Bigger Cells, Bigger Bees
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Assessment of Cell Size
The Cell Base
CHAPTER 10
The Construction of Combs
Parallelism between Combs
Festoons and Torsion
Festoons and Comb Growth
Evidence of a Sense of Equilibrium
Application of the Sense of Equilibrium
The Orientation of Combs
PART III: STIMULI FOR PRODUCTION AND MAN I PULATION OF WAX
CHAPTER 11
The Nectar Flow
The Stimulus of Spring
Winter, the Off-Switch
Nectar, the Unqualified Stimulus
Hoarding Assays
Comb as a Stimulus
The Honey Stomach
CHAPTER 12
Pollen and Wax Production
Essential role of pollen
Quantitative Requirements
Compartmental Effects
Physical Presence of Pollen
Pollen Pheromones
CHAPTER 13
The Brood Nest
The Meaning of Brood
Efficacy of Open Brood
Brood-Rearing and Honey Storage
CHAPTER 14
The Queen
The Construction of Queen Cells
Comb Construction and the Queen
XI
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CHAPTER 15
Space and Density
Nest Founding
Volume of a Potential Nest
The Arrangement of Space
Density Versus Space
Reduction of Nest Size
Quality of Space
REFERENCES
AUTHOR INDEX
SUBJECT INDEX
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