Fr. Michael Rennier 

How beekeeping satisfies the soul

Keeping bees is both a meaningful and sacred hobby … and deeply connected to the liturgy.

This past year, our family became apiarists. We got hold of a few colonies of honeybees and set them up in two hives side by side in front of a field of clover.

The bees have the run of 20 acres of a little wild and wooded valley tucked a few miles north of the Missouri River. They’re free to roam where they will, gathering nectar from wild clover, grape vines, and dogwood.

We’re not particularly talented beekeepers. One of the colonies seems far more motivated than the other – each takes its personality from their queen – and we don’t quite know how to fix it, but so far they’ve survived and even rewarded us with some of their extra honey.

Bees play a vital, if unseen, role in making the world beautiful. They have a complicated relationship with flowers, which tempt them in with bright, colored displays and sweet nectar only to secretly send them along with pollen stuck to their legs. I don’t think the bees are complaining, though.

Neither should we. After all, bees are the hidden laborers who, by spreading that pollen around, fertilize the flowers and make plants capable of producing fruit. Without bees, we wouldn’t know what an apple tastes like. Fruit would barely exist at all.

For example, the Central Valley in California, an area the size of Delaware, produces 2.3 billion pound of almonds every year, but those enormous orchards wouldn’t manage to produce even a fraction of those almonds without the help of bees. Beekeepers actually drive their colonies in on flatbed trucks for the week to help out.

Today it’s a widespread hobby and something of a big business, but it wasn’t long ago that beekeeping was a more specialized pursuit of monks and priests. Priests have always been interested in beekeeping – which is why I wanted to take it up also – not because of the insatiable desire for honey but because bees also make wax.

The reason we use beeswax candles for sacred liturgy

Beeswax makes the cleanest burning, brightest candles, which don’t produce a smell and don’t create a lot of smoke that leaves the ceiling and walls dirty with soot. Beeswax also burns longer than, say tallow candles made of animal fat.

Today, even though candles still burn on church altars every day, they’re an afterthought because we have electric lights. It used to be the case that candles not only provided an element of beauty and a comforting atmosphere, they were also vital to actually illuminating interior spaces.

To this very day, candles that burn on Catholic altars are required to be contain beeswax as the main ingredient.

There’s an interesting reason for this: Priests have always been aware that beeswax is a pure substance. Only the worker bees produce wax, and worker bees don’t mate with the queen. All their lives, they remain celibate and virginal. This is why candles – think, for instance, of the Paschal Candle – are symbols of Christ. They provide sacred light, as they burn the wax is consumed in sacrifice, and they’re made of virgin material.

The mystical quality of bees

Bees, it turns out, are highly theological. Maybe this is why they prompt endless poetic odes. Emily Dickinson, for instance, writes constantly about them;

The pedigree of honey
Does not concern the bee;
A clover, any time, to him
Is aristocracy.

As I watch our industrious little bees buzzing around their hives, I’m constantly amazed at their perseverance, the way they take pollen, a yellowish dust that most of despise and think of as the cause of sneezing misery, and use it to blanket the fields with flowers. Whatever pollen sticks to them, they turn into beauty.

We should all be more like bees. Dispensers of beauty. Or more like candles. A light cutting through shadows. These metaphors cling to me and I cannot help but hear in them the voice of God.

Perseverance, self-sacrifice, beauty, purity – all are habitual virtues I desire to practice with ever-greater dedication, and in this way transform my days into a continuous act of love. Each of us is a seed-sower of flowers in the fields.

Beeswax. What Is It?

The main constituents and the physical properties of beeswax.Wax is a hydrocarbon – contains the elements carbon, hydrogen and oxygen. It contains about 300 different chemicals.

Major components of a typical beeswax:

• Monohydric alcohols 31%
• Fatty acids 31%
• Hydrocarbons 16%
• Hydroxyl acids 13%
• Diols 3%
• Other (propolis, pollen, …) 6%

Physical properties of beeswax:

• Solid at room temperature
• Melts at 64°C
• Solidifies at 63°C
• It has a nice aroma (acids, alcohols, esters – volatiles)
• Specific gravity = 0.95. It floats in water and sinks in alcohol.
• It is not soluble in water.
• It is water repellent, used as water proofer – wax jackets and waxed threads.
• Slightly soluble in alcohol. Quite soluble in the higher order alcohols.
• It is soluble in chloroform, benzene, toluene, petrol, . . .
• It is brittle when cold.
• Malleable and plastic at 32°C
• It can remain stable for thousands of years.
• It is combustible, giving CO₂ and H₂O plus heat and light.

How Do I Make A Show Sample?

Purification.

• The wet cappings can be put in muslin bags (balanced) and spun in the extractor to recover as much honey as possible.
• I wash them in a little soft water to remove most of the honey and this can be used to feed some colonies – no waste.
• Rinse the cappings a few times in soft water. Hard water causes saponification with the calcium in the hard water – gives a bloom to the wax.
• For show the best wax comes from fresh cappings.
• Spread the cappings out on a white cloth and pick out any discoloured bits or foreign bodies – bee parts, propolis etc.
• The wax must be filtered through fine muslin. Place cappings in fine muslin cloth and tie it off.
• Place this in a jacketed hot water heater – when all of the wax is melted lift out the muslin slowly – reasonably pure wax will be left floating in the water.
• Filter this hot wax through fine filter paper to remove any further impurities, some use fine filter paper.
• Other methods can be used – solar extractor – steam boiler – or steam jacketed wax press.
• Do not use iron or copper vessels – stainlessless is preferred.

Mold

• Buy a mold of oven proof glass (Pyrex), free of internal blemishes and reserve for wax molding only.
• Check size of wax block from the show schedule.
• Put the appropriate volume of soft water in the mold and mark this level on the outside.
• Wash, with unscented detergent, and dry the mould.
• Heat the purified wax up to 70°C.
• Rub two to three drops soft water with two to three drops of unscented detergent all over the inside of the mold – to act as release agent.
• Place the mold in another larger Pyrex container with water at 66°C.
• Momentarily stand the heated wax container in a shallow tray of iced water – to congeal any dirt that may be in the bottom.
• Pour the molten wax into the mold, avoiding any air bubbles, up to the mark.
• Place a preheated lid on the mold.
• Allow to cool slowly, insulate with newspapers.
• Put the lot into the oven or the range when going to bed.
• The following morning submerge the mold in container of cold water – this releases the wax block.
• Smoothen off the edges and polish the surface with a silk cloth.
• The process will have to be repeated over several nights to get a block good enough for show exhibition.
• When you do get a good one – wrap it up well – put it in a plastic container and mind it.

reprinted with permission from

AN BEACHAIRE,

the  Irish Beekeeper

Ahhhh… Fall. This is the time of year that we finally get around to transforming our cappings into beautiful yellow blocks of wax. It is also the time that people start ordering candles again. Time to get busy making more candles.

So, what makes a great candle and how is it achieved? Beeswax candles are more than wax and a cotton string. They are a symbiotic relationship between air, wax, and wick. Since the most challenging candle to get right is the pillar candle, I will focus most of this article on the components of making a great pillar candle. In my opinion, the perfect pillar candle will create a burn pool that extends out most of the diameter of the candle, but not all the way. The flame is nice and bright with no smoke trails and the candle burns down all the way to the bottom without looking ugly and misshapen. The latter is really hard to achieve with taller candles so, although I make them, I prefer the small and medium sized candles for personal use.

I will start with wicks… Big sigh… This necessary part of the candle seems like it would be an easy thing, but honestly, although it can be maddening to figure out, I can’t stress enough how important this is to the overall quality of the candle. When a candle is lit, a series of events take place. First the match lights the wick and the wick itself starts to burn. The flame then starts to melt the wax. The wick acts as a pipeline that carries the melted wax in the form of a vapor to the flame via capillary action. Some wicks allow lots of fuel to flow quickly through a big pipe, while other wicks pump fuel more slowly through a smaller pipe. If you give the flame too much or too little fuel, it will burn poorly, or sputter out. The balance of fuel and flow needs to be just right.

Square Braid Wick Sizes – Wick graphic: Although this is not completely correct in terms of actual diameter of wicking, it gives an overall picture of relative sizes, commonly available wick sizes and the range of uses.

The nomenclature of square braid cotton wicking refers to the number of bundles, the ply of the wick, and how tightly it is braided. The 6/0 to 1/0 range of wicks, are constructed a bit differently than the larger wicks, but all of them are square which helps to channel the wax fumes up to the flame. It is important to keep your wicks well labeled and separated since similar sizes look identical. Often the only difference is the tightness of the braiding.

The design and nomenclature of this wicking is, I believe somewhat unique to the U.S. Apparently wicking produced and sold in other parts of the world utilize a different grading system and are not the same. The fact that what we have here in the US isn’t universal is something I discovered after my book was published and I was contacted by European editors that needed conversion factors for the wicking I recommended in my book.

Square Braid wick forms a carbon cap on the top of the wick. The carbon cap radiates heat outward from the flame which helps melt wax which is further away from the flame. The wick also bends slightly as it burns which minimizes carbon build-up and makes for a cleaner burning candle.

The oxygen seems like it would be the easy part- either the flame gets oxygen or it doesn’t. But the type of the candle and the environment that the candle is burned in play a role in how much oxygen the flame receives. I have found that the more open to air the flame is, the better the candle burned. So taper candles are perfectly set up for this. Pillars and votive candles, on the other hand typically start off burning beautifully, but as they burn wax and the flame travels downward into the candle, the flame often has problems. Either the flame tunnels down the middle of the candle melting very little wax and starving the flame of oxygen or the flame melts too much of the wax and flame is flooded and goes out.

#2/0 on the left and #2 on the right. As you can see, the actual diameter is about the same, bu the number of threads and the configuration of the thread clusters that make up the wicks are different.

So how does one ensure that the candle flame gets the oxygen it needs? Look at the burn pool. The width of the tunnel created by the burn pool is usually determined with the initial burning of the candle. The burn pool, which is the extent of melted wax, establishes the ultimate diameter of useable wax that the candle will ever use in subsequent burnings. The solid wax remaining around the outside of the burn pool will help the candle to retain its shape. For this reason, I always tell my customers that beeswax candles are intended to be burned all evening, not just a couple minutes and then extinguished. The combination of proper burning protocol and correct wick size should ensure that the burn pool reaches the desired width.

The blocks above, show the top and bottom of a wax block that I bought. From the top, it looks reasonably clean, but on the bottom, quite a bit of honey can be seen. This block will take some time and work before it can be used.

The last part of the candle trio is the beeswax itself. I personally hate to render wax, so I let my Karl handle the “heavy lifting” of rendering the cappings into big blocks of wax. I am not going to go into the rendering process here, since the process is often automated in larger operations. The wax that Karl renders out is really pretty clean, but since my candle business has outgrown what our hives can produce, I also purchase wax off another beekeeper in the area. His wax varies from relatively clean to blocks with rivers of honey buried inside.

For things like candles, especially pillar candles, the residual honey in the wax causes the wax to burn unevenly and to clog the wick. Even though a wax may “look” clean, it may still have honey in it.

The best way to get the last of the honey out of the wax is to allow it to clarify in a heated double boiler or wax tank. Admittedly, this task is easier to accomplish with the wax tank than a double boiler, since the wax needs to stay liquid for quite a while until all the honey has settled to the bottom. I usually let mine settle for a couple days. The best way to tell if it is done is by checking the clarity of the wax. When it is first melted, it has murkiness to it. As it settles, it starts to clarify. When the wax is clear, filter the wax through a clean piece of felt cloth and mold into useable portions. I usually do a variety of different sizes, so that I have the right size for whatever I am making. The resulting wax is still yellow and still has the signature honey-like scent, although the filtering may have lightened up the wax a little bit.

The dark spots are flecks of caramelized honey. Besides being unsightly, these are what ultimately can clog the wick, and ultimately keep the wax from reaching the flame.

The chart gives some general guidelines for wick sizes, but in order to ensure that the correct wick size is used, a burn test needs to be performed. Actually, the odds of getting it right on the first try is pretty rare, so this test probably needs to be performed multiple times until the right wick size is found.

The Right Size Wick
Use the following test to determine the proper wick size and scale up or down as needed.

Basic Burn Test
1. Trim the wick to a length of ¼” (6 mm). If you are testing more than one wick, make sure the candles are clearly labeled.

2. Place the test candles on a clean, flat, heat-resistant surface about 3” to 6” (7.5 cm to 15 cm) apart. Be sure to select a draft-free spot that is in full view of your workspace. Do not leave lit candles unattended.

3. Light the candles and record the time. It is critical to keep an eye on the candles while they are burning, especially when testing new wicks.

4. If testing pillar candles, allow them to burn for two hours then record the details of the melt pool and wick appearance. Ideally the melt pool will achieve the desired diameter by this point. If it hasn’t, the wick is most likely too small. Note any soot or mushrooming on the wick.

5. Allow the candle to burn for another four hours and record the details of the melt pool and wick again before gently blowing out the flame. At this point the melt pool of a well-wicked candle will have achieved the desired diameter and should be approximately ½” (1.3 cm) deep.

If the wick is mushrooming, the candle is sooting, or the melt pool is substantially deeper than ½” (1.3 cm), the wick is most likely too large.

6. Allow the candle to cool for at least five hours and repeat steps 4, 5, and 6 until the candle is completely burned. The quality of burn will almost always change during the entire burning of the candle. Burn the entire candle before deciding on a wick.

Wax from different batches can vary a bit not only in color, but also in behavior. Once the correct wick size is determined, test subsequent batches of wax to make sure that the candle still burns the way it should and if not, make the appropriate changes to the wick size.

So now, we can make some candles…

Petra Ahnert is a specialty candle designer living in the Milwaukee area, and is the author of Beeswax Alchemy.

by Clarence Collision

Wax is used by honey bees to protect themselves against water loss through the integument and in the construction of combs. The major fractions of the cuticular wax were analyzed by gas-liquid chromatography and were shown to be qualitatively similar to those of comb wax (Blomquist et al. 1980). However, the composition of the cuticular wax of the honey bee is quantitatively different from that of the comb wax. The major component of the cuticular lipids is hydrocarbon, which comprises 58% of this wax. In contrast, hydrocarbon comprises only 13-17% of the comb wax, and monoester is the largest component (Tulloch 1971). Comb wax is produced by four pairs of glands within the abdomen while cuticular wax is likely produced by epidermal cells of the integument.

Beeswax used in comb construction, comb repair, and capping of cells containing either pupae or honey, is secreted by worker bees on paired, smooth, oblong areas, called wax mirrors, located ventrally on abdominal segments four through seven (Figure 1A). On the dorsal side of each wax mirror is a layer of epithelial tissue called the wax gland (Sanford and Dietz 1976). Wax glands are merely specialized parts of the body-wall epidermis, which during the wax forming period in the life of the worker, become greatly thickened and take on a glandular structure (Figure 1C). The wax is discharged as a liquid through the mirrors and hardens to small flakes in the pockets between the mirrors and the long underlapping parts of the preceding sterna. After the wax-forming period the glands degenerate and become a flat layer of cells (Snodgrass and Erickson 1992).

The wax gland complex of the honey bee worker (Figure 2) consists of three cell types, epithelial cells, oenocytes and adipocytes (fat body cells), which act synergistically to secrete wax, a complex mixture of hydrocarbons, fatty acids and proteins (lipophorins) (Cassier and Lensky 1995). Lying over each gland is a large cellular mass composed of fat cells and oenocytes (Figure 1B) (Snodgrass 1956).
In young bees that have just recently emerged from their cells, the cells of the wax gland are nearly square in shape, while in older bees, the cells are either elongated or completely degenerated (Figure 1C). There is a definite correlation between the age of the bee, the size (length) of the cells in the wax glands, and the gland’s ability to secrete wax (Turell 1974). In newly emerged bees the wax gland cells have large nuclei and do not have intercellular spaces (an open area between the cells). These cubical cells have an average height of 17 to 19 microns. The production of wax begins in workers that are slightly less than one week old. As secretory activity increases, the cells of the wax gland become tall and slender. Cells from a bee at peak wax production have a height of approximately 50 microns, and have developed large intercellular spaces. Wax glands are best developed and most productive in 12-18 day-old workers. After producing wax for a few days, the wax glands begin to degenerate and by the time the bee is ready to leave the hive to become a field bee, usually when it is about 21 days of age, the glands have completely degenerated. The cellular boundaries, which were distinct in the immature and active glands, have become indistinct, or even lacking, and the height of the gland has fallen to only three microns.

Beeswax is produced by metabolizing honey in fat cells associated with the wax glands and converting it to beeswax; workers cannot produce beeswax unless there are adequate honey stores in the colony. Workers also need to eat pollen during the first five to six days of their life in order to secrete wax later on, evidently because the protein in pollen is needed at that time for adequate fat cell development (Winston 1987). Wax is secreted primarily during warm weather when foraging is active. Workers actively engaged in secreting wax engorge themselves with honey and hang in festoons at or near the site of comb building. Drones and queens do not have abdominal wax glands.

To date, the mechanisms associated with wax synthesis and secretion are not fully understood. Piek (1964) showed that acetic acid is likely taken up by the oenocytes and that acetate is used for the synthesis of hydrocarbons. Blomquist et al. (1980) demonstrated that the incorporation of injected radio-labelled acetate into hexane extractable wax (cuticular wax) by worker honey bees not actively producing comb wax resulted in the recovery of much of the radioactivity in the hydrocarbon fraction. In bees actively producing comb wax, a higher percentage of radioactivity was recovered in the monoester fraction. A dramatic effect of age on the distribution of radioactivity from acetate into the various wax fractions from bees studied in the Summer months was observed. In bees from 11 to 18 days following emergence to adults, the major wax component synthesized was monoester, whereas in younger and older bees, hydrocarbon was the major wax component formed. Both in vivo and in vitro experiments using bees actively producing comb wax showed that the abdomen produced significant amounts of monoester, hydrocarbon, and other esters, whereas the thorax synthesized mostly hydrocarbon. These data show that the epidermal cells and wax glands each produce a wax with a distinct composition, and that the age and seasonal differences observed in wax synthesis are due to presence or absence of active wax glands.

The ultra-structure of the cells of the wax gland complex was studied in relation to the synthesis and secretion of beeswax (Hepburn et al. 1991). The hydrocarbon and fatty acid profiles of epidermal cells and oenocytes were determined in relation to the ages of the bees. Smooth endoplasmic reticulum (SER)** was absent from both epidermal cells and adipocytes (fat cells) in adult workers from the time they emerged until the end of wax secretion. The oenocytes were rich in SER. The hydrocarbon and fatty acid content of the oenocytes, averaged for bee age, closely matched that of newly secreted wax. The oencytes are the probable source of the hydrocarbon fraction of beeswax which is consistent with histochemical and autoradiographic data. The cyclical changes of organelles within the cells and chemical composition of the wax gland complex closely coincided with measured, age-related rates of wax secretion in the workers.

Hepburn et al. (1991) observed that SER is barely discernible in the oenocytes of newly emerged workers, but by the fourth day the volume density of these tightly packed tubules is high.

Similarly, there is a large increase in whole oenocyte volume and the relative volumes of the oenocytes and SER remain elevated throughout the secretory phase. By day 18, both oenocytes and SER begin to decrease with the simultaneous appearance of primary lysosomes (membrane enclosed organelles in the cell that contain an array of digestive enzymes) and autolytic vacuoles (autolytic- cell destruction through the action of its own enzymes and vacuoles are organelles containing debris at various stages of degradation). Lipid and protein droplets were never observed in the oenocytes and cellular organelles showed no evident cyclical changes associated with wax synthesis. The fat cells are characterized by an extensive plasma membrane reticular system, numerous mitochondria (organelles that function in energy production), peroxisomes (organelles that contain enzymes involved in metabolic reactions) and rough endoplasmic reticulum and a few small Golgi bodies. The Golgi bodies or Golgi complex take up and process secretory products from the endoplasmic reticulum and then either releases the finished product into the cell cytoplasm or secretes them outside of the cell. SER is notably absent in fat cells from adult emergence through foraging. Massive lipid droplets occupy approximately 60% of cell’s cytoplasm in the young worker but decrease substantially over the next few days. Like the oencytes, the fat cells also increase in volume prior to wax synthesis. During wax synthesis, glycogen stores are notably large and the plasma membrane reticular system is well developed. As synthesis wanes, lipid droplets increase in size while the other organelles either remain unchanged or show small decreases in size. The notable and dynamic feature of the oenocyte is the abundant SER whose rise and fall are synchronized with measured periods of secretion (Hepburn et al. 1991). The major role of the epidermis in the production of wax appears to be the development of an elaborate system of small transport tubules (Locke 1961).

Figure 1.- The Wax Glands: A = Sternum of segment VI of worker, ventral, showing polished “mirrors” beneath wax glands, B = lengthwise section through two wax glands with overlying masses of fat cells and oenocytes. C = stages in the development and regression of a wax gland. Mir = mirror, WxGld = wax gland, FtCls = fat cells, Oen = oenocytes, vDph = ventral diaphragm, Mb = intersegmental membrane. (Snodgrass 1956).

The structure of the wax mirrors and the different types of cells were studied with scanning and transmission electron microscopy (Cassier and Lensky 1995). The outer face of the wax mirror shows a sub-regular, hexagonal pattern, each unit corresponding to an underlying epithelial cell. On perfectly clean wax mirrors, the covering epicuticle shows numerous holes or depressed areas from which the new biosynthesized wax masses exude, fuse and form irregular puddles. The sternal cuticle is particularly thin at the level of the mirror plates (two to four µm). It is composed of an outer trilaminate epicuticle, an homogeneous inner epicuticle and a two- or four-layered procuticle.

The main characteristic of the wax mirror is the presence of numerous pore canals containing microfilamentous structures or wax canal filaments (Locke 1961).

Beeswax when first secreted by the wax glands appears as a translucent white ellipsoidal flake. Production of beeswax in the honey bee is a process whereby many thin layers of wax are deposited on the wax mirror until a scale results. Dietz and Humphreys (1970) studied the structure of the wax scales with a scanning electron microscope. The customary shape of wax scales is due to the slightly recessed wax mirrors or plates which are situated below the wax glands. Essentially the wax scales produced by the wax glands of segments IV, V, and VI are somewhat similar in size and shape. Those originating with segment VII are not only smaller but also of different shape.

The mechanism by which beeswax penetrates the cuticle to form the layers of the scale has been subject of several investigations (Sanford and Dietz 1976). The extensive system of pore canals filled with filaments, believed to consist of wax extending up into the wax canals, is thought to form part of the transport mechanism which brings wax or its precursors near to the surface of the wax mirrors (Goodman 2003).

The cuticle of the wax mirrors has a stratified structure and the associated fibrous structures and pore canals certainly play a major function during the transit of the wax components from the wax glands to the exterior surface (Locke 1961). It has been hypothesized that the wax components are transported in a protein medium through the gland cells to the outer surface of the mirrors, where the molecules condense to form scales. Kurstjens et al. (1990) electrophoretically detected proteins in the wax scales and in the comb wax.

Cassier and Lensky (1995) with scanning and transmission electron microscopy showed the extrusion of wax droplets through the wax mirrors and for the first time large cisternae (fluid containing sac or cavity) of smooth endoplasmic reticulum in the epidermis. These cisternae are probably involved in the transit of wax from the oenocytes to the pore canal system. The cisternae can also convey apolipophorins from the hemolymph to the wax mirrors.

The elongate epithelial cells of the wax gland form a palisade layer. Intercellular spaces and infoldings of the plasma membrane delimit large spaces where twisted filamentous structures run and connect to those of the pore canals (Cassier and Lensky 1995). Beeswax is a composite mixture of hydrocarbons, esters, fatty acids (Hepburn et al. 1991) and proteins (Kurstjens et al. 1990). Because beeswax is hydrophobic it is probably transported from the wax glands to the mirrors by hemolymph lipophorins via the SER cisternae.

Based on biochemical and structural data, it is possible to suggest the contributions of each cell type to the wax gland complex. Oenocytes are involved in the secretion of the hydrocarbon fraction of the wax (Piek 1964; Blomquist and Ries 1979; Lambremont and Wykle 1979; Hepburn et al. 1991). The epithelial cell provided with ribosomes, polysomes, rough endoplasmic reticulum cisternae and electron-dense granules probably synthesizes a part of the protein fraction of the wax product, the other part coming directly from hemolymph through SER cisternae. Fat body cells provide plastic and energetic products. In contrast to previous workers, Cassier and Lensky (1995) found that the SER is well developed both in oenocytes and epidermal cells. In the epidermal cells, SER forms long cisternae parallel to the major axis, from the basal to the distal pole of the epithelial cells. They seem to be connected to extracellular spaces, thus forming a complete pathway to the wax mirror exterior.

Examination of the fine structure of the wax gland indicates that its function in beeswax secretion is either strictly transport or concentration of substances rather than producing them for release as in the case with many secretory cells (Sanford and Dietz 1976). Piek (1964) showed that the constituents of beeswax are synthesized in the fat cells and oenocytes, which are enzyme activated by an esterase, which could possibly catalyze wax production in the cuticle as well as in the epithelium. Piek concluded that esters are synthesized by fat cells and hydrocarbons and wax acids by the oenocytes. These wax precursors are then discharged into the wax gland (epithelium), concentrated and then pumped into the extracellular space.

The actual site of the final reactions that complete beeswax formation before it is secreted through the pore canals is not known. It may be assumed from its accumulation as a liquid layer on the wax mirrors, and from its solidification rapidly after secretion, that the final reactions that result in hardening of the wax take place after secretion.

Beewax is a complex substance made up of wax esters, fatty acids and hydrocarbons (Piek 1964; Tulloch 1970). Over 300 individual chemical components have been identified from pure beeswax (Tulloch 1980). Beeswax consists primarily of monoesters (35%), hydrocarbons (14%), diesters (14%), triesters (3%), hydroxymonoesters (4%), hydroxypolyesters (8%), free fatty acids (12%), acid esters (1%), acid polyesters (2%), free alcohol (1%) and unidentified (6%). It is this great diversity of composition that gives beeswax many unique properties (Goodman 2003) and keeps us from fully understanding the synthesis and secretion process.

References

Blomquist, G.J. and M.K. Ries 1979. The enzymatic synthesis of wax esters by a microsomal preparation from the honey bee Apis mellifera L. Insect Biochem. 9: 183-188.
Blomquist, G.J., A.J. Chu and S. Remaley 1980. Biosynthesis of wax in the honeybee, Apis mellifera L. Insect Biochem. 10: 313-321.
Cassier, P. and Y. Lensky 1995. Ultrastructure of the wax gland complex and secretion of beeswax in the worker honey bee Apis mellifera L. Apidologie 26: 17-26.
Dietz, A. and W.J. Humphreys 1970. Scanning electron microscopy study of the structure of honey bee wax scales. J. Georgia Entomol. Soc. 5: 1-6.
Goodman, L. 2003. Form and Function in the Honey Bee. International Bee Research Association, Cardiff, UK, 220 pp.
Hepburn, H.R., R.T.F. Bernard, B.C. Davidson, W.J. Muller, P. Lloyd, S.P. Kurstjens and S.L. Vincent 1991. Synthesis and secretion of beeswax in honey bees. Apidologie 22: 21-36.
Kurstjens, S.P., E. McClain and H.R. Hepburn 1990. The proteins of beeswax. Naturwissenschaften 77: 34-35.
Lambremont, E.M. and R.L. Wykle 1979. Wax synthesis by an enzyme system from the honey bee. Comp. Biochem. Physiol. 63B: 131-135.
Locke, M. 1961. Pore canals and related structures in insect cuticle. J. Biophys. Biochem. Cytol. 10: 589-618.
Piek, T. 1964. Synthesis of wax in the honeybee (Apis mellifera L.) J. Insect Physiol. 10: 563-572.
Sanford, M.T. and A. Dietz 1976. The fine structure of the wax gland of the honey bee (Apis mellifera L.). Apidologie 7: 197-207.
Snodgrass, R.E. 1956. Anatomy Of The Honey Bee, Comstock Publ. Assoc., Ithaca, NY, 2nd ed.
Snodgrass, R.E. and E.H. Erickson 1992. The anatomy of the honey bee. In The Hive And The Honey Bee, Dadant and Sons, Inc., Hamilton, IL. pp. 103-169.
Tulloch, A.P. 1970. The composition of beeswax and other waxes secreted by insects. Lipids 5: 247-258.
Tulloch, A.P. 1971. Beeswax: structure of esters and their component hydroxyl acids and diols. Chemy. Phys. Lipids. 6: 235-265.
Tulloch, A.P. 1980. Beeswax- composition and analysis. Bee Wld. 61: 47-62.
Turell, M.J. 1974. The wax glands of the honey bee. Am. Bee J. 114: 328-330.
Winston, M.L. 1987. The Biology Of The Honey Bee. Harvard University Press, Cambridge, MA, 281 pp.

Clarence Collison is an Emeritus Professor of Entomology and Department Head Emeritus of Entomology and Plant Pathology at Mississippi State University, Mississippi State, MS.

** Endoplasmic reticulum is a membrane system of folded sacs and interconnected channels within the cell’s cytoplasm that serve as a site for protein and lipid (fat) synthesis. Rough endoplasmic reticulum (RER) is in the form of flat bands covered with ribosomes that are responsible for the synthesis of many proteins. Smooth endoplasmic reticulum (SER) is tubular in form; serving as the site of lipid synthesis and carbohydrate metabolism.