Module 5 Study Notes

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Module 5 Study NotesIntroductionReferences:The Honeybee Inside OutThe Honeybee Around and AboutGuide to Bees and HoneyBeekeeping Study Notes (modules 5,6,7&8)The Biology of the Honey BeeThe Anatomy of the Honey BeeCelia F. DavisCelia F. DavisTed HooperJ.D & B.D. YatesMark L. WinstonR. D. SnodgrassBBKA websiteMBBKA Study GroupMBBKA Basic Course Notes21/1/2011Page 1

Module 5 Study NotesContentsThe Candidate shall be able to describe in detail and illustrate where appropriate, referring tohistological features as necessary:5.1 the alimentary system including the process of digestion by enzymes and the absorption andassimilation of products of digestion; . 35.2 the excretory system and the substances excreted; . 55.3 the respiratory system, including the muscular ventilation of the air sacs, the structure andoperation of the spiracles and exchange of respiratory gases; . 75.4 the circulatory system, including the heart, dorsal and ventral diaphragms and the compositionand functions of the haemolymph; . 115.5The exocrine glands of all castes and sexes of adult bees and larvae, the functions and maincompositions of their secretions including pheromones, (hypopharyngeal, mandibular, tergite glandsof the queen (Renner-Baumann); Nasonov, sting, Arnhart post cerebal, thoracic salivary, wax glandsand wax production) . 125.6 the structure and function of the nervous system and sense organs (including the compound eyes,ocelli, organ of Johnston and the sensilla); . 155.7 the endocrine glands and the functions of their secretions particularly the neurosecretary cells, thecorpora allata, corpora cardiac and the prothoracic glands; . 185.8 the fat body and its storage of metabolites; . 205.9 the reproductive system of queen and drone and the production of sperm and eggs; . 215.10 the structure of the egg, development of the embryo within the egg and the hatching of the larva;. 245.11 the external and internal structure of the honeybee larva;. 255.12 the metamorphosis of the larva with outline accounts of ecdysis, larval defaecation, cocoonspinning, the external anatomy of the pro-pupa, its change to a pupa and then to an imago; . 275.13 the effect of feeding and other factors on caste determination including discussion about thedifferences between brood food and royal jelly;. 295.14 the physiological and structural differences between laying workers and normal workers and therole of pheromones in bringing about these differences; . 315.15 the differences between summer and winter worker honeybees; . 325.16 the structure and main constituents of the cuticle with an outline account of its invagination withinthe body to form linings of the gut and tracheae; . 335.17 the external anatomy of the queen, worker and drone; . 355.18 the function and structure of the wings, legs, feet, antennae, mouth parts and setae (hairs); . 375.19 the structure of the sting mechanism and how this mechanism operates to penetrate human skinand deliver the venom; . 445.20 the role of the direct and indirect muscles in flight. . 4721/1/2011Page 2

Module 5 Study Notes5.1 the alimentary system including the process of digestion by enzymes andthe absorption and assimilation of products of digestion;The digestive system of any animal has four distinct functions Ingestion – taking in of foodDigestion – breaking the food down chemically to simplest, soluble componentsAbsorption – passing those components through its wall to the bloodDefecation – removing any indigestible material from the bodyThe honey bee is more complicated due to the need to transport nectar and water.Food is broken down by the process of digestion and these products are then circulated by thehymolympth (blood) and used to provide energy, bodybuilding substances, and the requirements forcarrying out the chemical processes of life. The waste products of these processes have to becollected and eliminated from the insect’s body. Digestion and excretion are the functions of thealimentary canal and its associated glands. These are shown in the figure. The mouth is between thebase of the mandibles below the labrum and above the labium. Immediately inside the mouth thecanal expands into a cavity, formed by the cibarium and pharynx, which has muscular attachments tothe front of the head which can expand and contract it, thus providing small amounts of suction tohelp pass the food from the proboscis and into the oesophagus.Two pairs of glands empty into the cibarium: Postcerebral glandsThoracic glandsThese glands are collectively called labial or salivary glands. They produce a liquid that passes underthe pharynx and opens into the base of the labium and on to the proboscis where it is mixed withincoming food. The saliva contains invertase (from the hypopharyngeal glands) which begins thebreakdown of sucrose.Muscles in the oesophagus provide waves of contractions which work the food back into the dilatedcrop or “honey stomach”, where it is stored for a while. At the end of the honey stomach is the21/1/2011Page 3

Module 5 Study Notesproventriculus, a valve that prevents the nectar from going any further unless it needs some for itsown use.If the bee is a forager it is here that the nectar is carried back to the colony and regurgitated back tothe mouth and fed to other bees. The proventriculus has four lips which are in continuous movement,sieving out solids from the nectar. The solids – pollen grains, spores, even bacteria – are removedfrom the nectar fairly quickly and passed back as a fairly dry lump, or bolus, into the ventriculus.When the bee needs to have sugar the whole proventriculus gapes open and an amount of nectar isallowed through to the ventriculus, where the food is subjected to several enzymes which break itdown into molecules small enough to be passed through the gut wall into the hymolympth. The beeappears to only digest two main food types, sugars and proteins. These are digested by enzymesproduced in the walls of the ventriculus, assimulated and used to produce energy or to build up thebees own proteins.The vetriculus is called the mid gut or true stomach, it is lined with epithelium which releases enzymesin order to breakdown fats, proteins and complex sugars into simpler constituent parts. The food isencompassed by a peritophic membrane which again is produced by the epithelium, this film protectsthe thin wall of the ventriculus during the food passage through it. Non-soluble products pass throughto the small intestine.The small intestine is again lined with epithelium to enable further digestion. The residue is passedinto the rectum where it is held, as faeces, until the bee is able to leave the hive and void the contentsof the rectum in flight. During long spells of cold weather in winter the rectum can extend almost thewhole length of the abdomen before the bee is able to get out for a cleansing flight. At the end of theventriculus are about a hundred small thin walled tubes. These are the malpighian tubules which havea similar function to our kidneys in that they remove nitrogenous waste (the results of the breakdownof proteins during metabolism) from the hymolympth. The waste products mainly in the form of uricacid are passed into the gut to join the faeces in the rectum.21/1/2011Page 4

Module 5 Study Notes5.2 the excretory system and the substances excreted;The excretory system is essentially a sophisticated filtration system which not only removes wastesubstances, which would slowly poison the cells, but also acts selectively, adjusting the amounts ofparticular substances in the haemolymph so that there is a balance between water and salts and theosmotic pressure and acidity remain within narrow limits.There are two types of waste produced by active cells: Carbon Dioxide produced as a result of respiration and removed by the respiratory systemNitrogenous waste resulting from the chemical reactions which go on within the cells involvingproteins and other nitrogen-containing compounds, this waste is removed by the excretorysystem.There are four aspects of the excretory system; Filtration of the haemolymph by the Malpighian tubulesRe-absorption from the excretory system of useful substancesActive secretion of substances into the systemComplete removal of the end product to the outside of the bees bodySubstances in the haemolymph are filtered through the wall of the Malpighian tubule at its upper(distal) end. Muscle fibres cause the tubule to wave about and come in contact with the maximumamount of haemolymph. The substance passes through the single cell wall and travels down thecentre (lumin) of the tubule.As materials travel down the lumin some are re-absorbed, water retention is crucial part of reabsorbtion, depending on the Haemolymph state salts may or may not be re-absorbed.Finally, other substances are actively secreted by cells. They pull passing molecules in and pushthem through into the lumin. The Malpighian activity requires a lot of energy hence the proximity oftracheoles for sourcing Oxygen.When the material in the lumen of the tubule reaches the intestinal connection, it passes into thedigestive system and together with the waste from the digestive system is passed to the outside of the21/1/2011Page 5

Module 5 Study Notesbody through the small intestine, the rectum and finally the anus. During its final passage furtherwater is re-absorbed.21/1/2011Page 6

Module 5 Study Notes5.3 the respiratory system, including the muscular ventilation of the air sacs,the structure and operation of the spiracles and exchange of respiratorygases;SummaryThe bee breathes through tubes called trachea which convey oxygen to where it is required within thebody of the insect. In all the higher animals oxygen is transported to the tissues by blood, bit in insectsthe blood is not involved in the transport of oxygen through the body. The trachea is made of cuticleand is prevented from collapsing by spiral thickening. The trachea start quite large but very rapidlydivide many times, getting smaller all the while, until finally they end up as single cells, or a loop. Thetrachea open to the air through holes in the cuticle called spiracles, and in many cases these areprovided with a closing mechanism.Air enters the tracheal system through spiracles and fills the tubes. When the cells in which thetrachea end are using up oxygen, this reduces the pressure of oxygen at that point and molecules ofoxygen migrate in to make up the deficiency. It is thus by diffusion that oxygen makes its way via thetrachea into the body of the bee. Oxygen is used to oxidize substances such as sugar in the cells torelease energy for their use, producing the residue substances, carbon dioxide and water, this iscellular respiration. In the honeybee the main tracheal trunks become large sacs which are ventilatedby breathing movements of the abdomen, whereby the abdomen is lengthened and contracted in atelescopic type movement.21/1/2011Page 7

Module 5 Study NotesDefinitions:The respiratory system comprises Spiracles (openings), Trachea (large tubes), tracheoles (smallertubes) and the total surface through which oxygen and carbon dioxide is exchanged is called therespiratory surface. The respiratory surface must be moist for the gases to diffuse, the tracheolestherefore contain fluid.SpiraclesThere are 10 pairs, found on each segment from T2 through A8, There is usually a valve surroundedby hairs on each spiracle, exceptions being the small spiracles on T3 and the “lid” on T2 not beingable to fully close. The two spiracles on the propodeum A1 are the largest and all the Abdominalspiracles have a widened entrance called an Atrium. The valves are controlled by small muscles.TracheaeThe tracheae are tubes leading from the spiracles and joining together in the body to give two largelateral tracheae (air sacs in adult bee). The tracheae have thicker areas (made of chitin) like a spiralaround the tube which prevent it from collapsing in low pressure. The lining of the tracheae is cuticleand is contiguous with the outer cuticle of the bee (implying it sheds with each molt).21/1/2011Page 8

Module 5 Study NotesTracheolesThe tracheae split down to smaller tubes (traceoles) which terminate in very small blind endings withfluid for the diffusion of gases. Whether the end is a single cell, disturbs or splits cells is not known,but they will generally terminate where needed such as close to muscle tissue.Air SacsAre present in the head, thorax and abdomen of the bee, they are tracheae with no thickening(taenidia) so have thin walls and can expand and contract. This is enables the bee to breath:Build up of carbon dioxide stimulates the nerves in the ganglia which control the muscles thatstretch and contract the membrane linking the sternum and tergum. Build up of carbondioxide contracts the muscles forcing it out via the spiracles, releasing the muscles causes avacuum allowing fresh air in. A similar action occurs longitudinally in the abdomen as well asin the thorax.21/1/2011Page 9

Module 5 Study NotesBreathingWhen resting or working most air flows in and out through the spiracles on T2, when flying whichrequires large amounts of oxygen, air mainly flows in T2 and out through spiracles on A1.21/1/2011Page 10

Module 5 Study Notes5.4 the circulatory system, including the heart, dorsal and ventral diaphragmsand the composition and functions of the haemolymph;The circulatory system is an Open System, where fluid (Haemolymph) flows throughout the bodysupporting cells, transporting food and removing the waste products of metabolism.The elements that control the fluid and effectively comprise the circulatory system are:-HeartAortaAntennal vesticleDorsal diaphragmVentral diaphragmThe heart is tube which is closed at the rear end and has 5 pairs of openings along its length calledostia, haemolymph is drawn into these openings and pumped forward to the aorta. The heart resideswithin the Dorsal sinus and is connected to the body and the dorsal diaphragm.The aorta is a narrow tubular extension to the front of the heart which passes through the petiole andopens out at the brain.At the base of the antennae there is a structure that draws in haemoplymph when relaxed and pumpsit up the antennae when tensioned.The dorsal diaphragm is connected in places to the upper part of the abdomen, above the diaphragmis a gap (sinus) within which the heart resides. The diaphragm has rhythmic undulations from the rearof the abdomen to the front which causes surrounding fluid to travel forwards.The ventral diaphragm exists within the thorax and the abdomen and is attached to the sternum,again there is a sinus between the diaphragm and the body of the bee. The rhythmic movement of theventral diaphragm is front to back, moving fluid from thorax to abdomen within the sinus.The fluid within the space between the diaphragms swirls.The haemolymph comprises; plasma and haemocytes. Plasma is a colourless liquid (90% water)containing several dissolved substances including salts, amino acids, proteins, carbohydrates, uricacid, lipids, fatty acids and organic compounds. The haemocytes are simple cells suspended in theplasma and are instrumental in coagulation and wound healing.The functions of Haemolymph can be described as:Transport, of food to cells, waste materials after metabolism, hormones from organ source to point ofuseMechanical support, it fills the cavities of the beeControl of water content of the cells, makes water and dissolved substances available to the cellsMetabolism, chemicals are broken down into simpler ones in the haemolymph, essential for creationof energyPhagocytosis, some of the haemocytes destroy bacteria and parasitesWound healing, haemocytes form chain to plug holes until wound repairs.21/1/2011Page 11

Module 5 Study Notes5.5The exocrine glands of all castes and sexes of adult bees and larvae, the functions and main compositions of theirsecretions including pheromones, (hypopharyngeal, mandibular, tergite glands of the queen (Renner-Baumann);Nasonov, sting, Arnhart post cerebal, thoracic salivary, wax glands and wax nt of headAbovemandiblesFunctionProduces element of brood food inyounger worker and enzymes in olderworkerGorging on pollen can cause gland torevert to food productionYoung worker, production of brood foodand royal jellyMature worker, alarm pheromone issuedby guard bees to ward off robbers andinitiate stinging response from other beesTergite (rennerBaumann)Edges ofabdominaltergites A3-5NasonovTergite A721/1/2011Queen, produces pheromones used inmating or part of queen substanceQueen recognition, contributes to QueenSubstance, emitted through Queengrooming and retenue palpating herabdomen with their antennaeLocation scent used when flying in swarmto attract other bees, marking theentrance to hive, marking source of waterCompositionYoung worker protein in form of clear liquid for making broodfoodSucrase and glucose oxidase in older/foraging workerCasteWIn young worker 10-HDO (10-hydroxydec-2-enoic acid) whichis principle fatty acid in brood food and acts as preservativeOlder worker 2-heptanone which is the alarm pheromoneQWDQueen produces :9-oxodec-2-enoic acid (drone attractant when mating)9-hydroxydec-2-enoic acid (holds swarm together)Composition unknownTerpenic alcoholsTerpenic aldehydesTerpenic acidsQGeraniol, Nerol and (E,E)-Farnesol(E)-Citral and (Z)-CitralGeranic and Nerolic acidWPage 12

Module 5 Study NotesSting scentglandQuadrate platesSting acid(poison orverom gland)Sting Alkaline(Dufour)Abdomenpost cerebalBehind brainSting ChamberThoracicIn the thoraxArnhart(tarsal glands)5 tarsome ofeach legthWax glandsSternites A4-7Wax productionSternites A4-721/1/2011Alarm pheromone, (Z). attracts bees tosting site and stabilises Isopentyl whichalong with 2-heptanone elicits stingingresponseProduction of venom to be used in sting(Z)-11-Eicosen-1-olIsopentyl acetateWMajor ones: Mellitin (50% of dry weight), Phospholipase A,Hyaluronidase, Acid phosphatise, Allergen CQWGenerally unknown but assumed to:lubricate sting mechanism, neutraliseremaining acid and in queen protectivecoating to eggs or egg adhesive for cellfloorSalivary, no reservoir, secrete directly intooutlet ductsUnknown, white in colour and alkaline in natureQWWater of Alkaline natureQWDWater of Alkaline natureQWDUnknown except different in the two castesQWOnly vestigial in drone, equaldevelopment in Queen and W

Module 5 Study Notes 21/1/2011 Page 1 Introduction References: The Honeybee Inside Out Celia F. Davis The Honeybee Around and About Celia F. Davis Guide to Bees and Honey Ted Hooper Beekeeping Study Notes (modules 5,6,7&8) J.D & B.D. Yates The Biology of the Honey Bee Mark L. Winston The Anatomy of the Honey Bee R. D. Snodgrass BBKA website MBBKA Study Group MBBKA Basic Course Notes . Module 5 .

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