By J. W. WHITE, JR. AND LANDIS W. DONER(1)
BEEKEEPING IN THE UNITED STATES
AGRICULTURE HANDBOOK NUMBER 335
Revised October 1980
Pages 82 - 91

Honey is essentially a highly concentrated water solution of two sugars, dextrose and levulose, with small amounts of at least 22 other more complex sugars. Many other substances also occur in honey, but the sugars are by far the major components. The principal physical characteristics and behavior of honey are due to its sugars, but the minor constituents - such as flavoring materials, pigments, acids, and minerals - are largely responsible for the differences among individual honey types.

Honey, as it is found in the hive, is a truly remarkable material, elaborated by bees with floral nectar, and less often with honeydew. Nectar is a thin, easily spoiled sweet liquid that is changed (”ripened”) by the honey bee to a stable, high-density, high-energy food. The earlier U.S. Food and Drug Act defined honey as “the nectar and saccharine exudation of plants, gathered, modified, and stored in the comb by honey bees (Apis mellifera and A. dorsata); is levorotatory; contains not more than 25% water, not more than 0.25% ash, and not more than 8% sucrose.” The limits established in this definition were largely based on a survey published in 1908. Today, this definition has an advisory status only, but is not totally correct, as it allows too high a content of water and sucrose, is too low in ash, and makes no mention of honeydew.

Colors of honey form a continuous range from very pale yellow through ambers to a darkish red amber to nearly black. The variations are almost entirely due to the plant source of the honey, although climate may modify the color somewhat through the darkening action of heat.

The flavor and aroma of honey vary even more than the color. Although there seems to be a characteristic “honey flavor,” almost an infinite number of aroma and flavor variations can exist. As with color, the variations appear to be governed by the floral source. In general, light-colored honey is mild in flavor and a darker honey has a more pronounced flavor. Exceptions to the rule sometimes endow a light honey with very definite specific flavors. Since flavor and aroma judgments are personal, individual preference will vary, but with the tremendous variety available, everyone should be able to find a favorite honey.

—-
(1) Research leader and research chemist, respectively, Science and Education Administration, Eastern Regional Research Center, Philadelphia, Pa. 19118.

Composition of Honey

By far, the largest portion of the dry matter in honey consists of the sugars. This very concentrated solution of several sugars results in the characteristic physical properties of honey - high viscosity, “stickiness,” high density, granulation tendencies, tendency to absorb moisture from the air, and immunity from some types of spoilage. Because of its unique character and its considerable difference from other sweeteners, chemists have long been interested in its composition and food technologists sometimes have been frustrated in attempts to include honey in prepared food formulas or products. Limitations of methods available to earlier researchers made their results only approximate in regard to the true sugar composition of honey. Although recent research has greatly improved analytical procedures for sugars, even now some compromises are required to make possible accurate analysis of large numbers of honey samples for sugars.

An analytical survey of U.S. honey is reported in Composition of American Honeys, Technical Bulletin 1261, published by the U.S. Department of Agriculture in 1962. In this survey, considerable effort was made to obtain honey samples from all over the United States and to include enough samples of the commercially significant floral types that the results, averaged by floral type, would be useful to the beekeeper and packer and also to the food technologist. In addition to providing tables of composition of U.S. honeys, some general conclusions were reached in the bulletin on various factors affected by honey composition.

Where comparisons were made of the composition of the same types of honey from 2 crop years, relatively small or no differences were found. The same was true for the same type of honey from various locations. As previously known, dark honey is higher than light honey in ash (mineral) and nitrogen content. Averaging results by regions showed that eastern and southern honeys were darker than average, whereas north-central and intermountain honeys were lighter. The north-central honey was higher than average in moisture, and the intermountain honey was more heavy bodied. Honey from the South Atlantic States showed the least tendency to granulate, whereas the intermountain honey had the greatest tendency.

The technical bulletin includes complete analyses of 490 samples of U.S. floral honey and 14 samples of honeydew honey gathered from 47 of the 50 States and representing 82 “single” floral types and 93 blends of “known” composition. For the more common honey types, many samples were available and averages were calculated by computer for many floral types and plant families. Also given in this bulletin are the average honey composition for each State and region and detailed discussions of the effects of crop year, storage, area of production, granulation, and color on composition. Some of the tabular data are included in this handbook.

Table 1 gives the average value for all of the constituents analyzed in the survey and also lists the range of values for each constitutent. The range shows the great variability for all honey constituents. Most of the constituents listed are familiar. Levulose and dextrose are the simple sugars making up most of the honey. Fructose and glucose are other commonly used names for these sugars. Sucrose (table sugar) also is present in honey, and is one of the main sugars in nectar, along with levulose and dextrose. “Maltose” is actually a mixture of several complex sugars, which are analyzed collectively and reported as maltose. Higher sugars is a more descriptive term for the material formerly called honey dextrin.

The undetermined value is found by adding all the sugar percentages to the moisture value and subtracting from 100. The active acidity of a material is expressed as pH; the larger the number the lower is the active acidity. The lactone is a newly found component of honey. Lactones may be considered to be a reserve acidity, since by chemically adding water to them (hydrolysis) an acid is formed. The ash is, of course, the material remaining after the honey is burned and represents mineral matter. The nitrogen is a measure of the protein material, including the enzymes, and diastase is a specific starch-digesting enzyme.

Most of these constituents are expressed in percent, that is, parts per hundred of honey. The acidity is reported differently. In earlier times, acidity was reported as percent formic acid. We now know that there are many acids in honey, with formic acid being one of the least important. Since a sugar acid, gluconic acid, has been found to be the principal one in honey, these results could be expressed as “percent gluconic acid” by multiplying the numbers in the table by 0.0196. Since actually there are many acids in honey, the term “milliequivalents per kilogram” is used to avoid implying that only one acid is found in honey. This figure is such that it properly expresses the acidity of a honey sample independently of the kind or kinds of acids present.

In table 1, the differences between floral honey and honeydew honey(2) can be seen. Floral honey is higher in simple sugars (levulose and dextrose), lower in disaccharides and higher sugars (dextrins), and contains much less acid. The higher amount of mineral salts (ash) in honeydew gives it a less active acidity (higher PH). The nitrogen content reflecting the amino acids and protein content is also higher in honeydew.

The main sugars in the common types of honey are shown in table 2. Levulose is the major sugar in all the samples, but there are a few types, not on the list, that contain more dextrose than levulose (dandelion and the blue curls). This excess of levulose over dextrose is one way that honey differs from commercial invert sugar. Even though honey has less dextrose than levulose, it is dextrose that crystallizes when honey granulates, because it is less soluble in water than is levulose. Even though honey contains an active sucrose-splitting enzyme, the sucrose level in honey never reaches zero.

Honey varies tremendously in color and flavor, depending largely on its floral source. Its composition also varies widely, depending on its floral sources (table 2). Although hundreds of kinds of honey are produced in this country, only about 25 or 30 are commercially important and available in large quantities. Until the comprehensive survey of honey composition was published in 1962, the degree of compositional variation was not known. This lack of information hindered the widespread use of honey by the food industry.

Water Content

The natural moisture of honey in the comb is that remaining from the nectar after ripening. The amount of moisture is a function of the factors involved in ripening, including weather conditions and original moisture of the nectar. After extraction of the honey, its moisture content may change, depending on conditions of storage. It is one of the most important characteristics of honey influencing keeping quality, granulation, and body.

Beekeepers as well as honey buyers know that the water content of honey varies greatly. It may range between 13 and 25 percent. According to the United States Standards for Grades of Extracted Honey, honey may not contain more than 18.6 percent moisture to qualify for U.S. grade A (U.S. Fancy) and U.S. grade B (U.S. Choice). Grade C (U.S. Standard) honey may contain up to 20 percent water; any higher amount places a honey in U.S. grade D (Substandard).

These values represent limits and do not indicate the preferred or proper moisture content for honey. If honey has more than 17 percent moisture and contains a sufficient number of yeast spores, it will ferment. Such honey should be pasteurized, that is, heated sufficiently to kill such organisms. This is particularly important if the honey is to be “creamed” or granulated, since this process results in a slightly higher moisture level in the liquid part. On the other hand, it is possible for honey to be too low in moisture from some points of view. In the West, honey may have a moisture content as low as 13 to 14 percent. Such honey is somewhat difficult to handle, though it is most useful in blending to reduce moisture content. It contains over 6 percent more honey solids than a product of 18.6 percent moisture.

In the 490 samples of honey analyzed in the Department’s Technical Bulletin 1261, the average moisture content was 17.2 percent. Samples ranged between 13.4 and 22.9 percent, and the standard deviation was 1.46. This means that 68 percent of the samples (or of all U.S. honey) will fall within the limits of 17.2 ± 1.46 percent moisture (15.7 - 18.7); 95.5 percent of all U.S. honey will fall within the limits of 17.2 ± 2.92 percent moisture (14.3 - 20.1).

In the same bulletin, a breakdown of average moisture contents by geographic regions is shown. These values (percent) are North Atlantic, 17.3; East North Central, 18.0; West North Central, 18.2; South Atlantic, 17.7; South Central, 17.5; Intermountain West, 16.0; and West, 16.1.

Sugars

Honey is above all a carbohydrate material, with 95 to 99.9 percent of the solids being sugars, and the identity of these sugars has been studied for many years. Sugars are classified according to their size or the complexity of the molecules of which they are made. Dextrose (glucose) and levulose (fructose), the main sugars in honey, are simple sugars, or monosaccharides, and are the building blocks for the more complex honey sugars. Dextrose and levulose account for about 85 percent of the solids in honey.

Until the middle of this century, the sugars of honey were thought to be a simple mixture of dextrose, levulose, sucrose (table sugar), and an ill-defined carbohydrate material called “honey dextrin.” With the advent of new methods for separating and analyzing sugars, workers in Europe, the United States, and Japan have identified many sugars in honey after separating them from the complex honey mixture. This task has been accomplished using a variety of physical and chemical methods.

Dextrose and levulose are still by far the major sugars in honey, but 22 others have been found. All of these sugars are more complex than the monosaccharides, dextrose and levulose. Ten disaccharides have been identified: sucrose, maltose, isomaltose, maltulose, nigerose, turanose, kojibiose, laminaribiose, a, B-trehalose, and gentiobiose. Ten trisaccharides are present: melezitose, 3-a-isomaltosylglucose, maltotriose, l-kestose, panose, isomaltotriose, erlose, theanderose, centose, and isopanose. Two more complex sugars, isomaltotetraose and isomaltopentaose, have been identified. Most of these sugars are present in quite small quantities.

Most of these sugars do not occur in nectar, but are formed either as a result of enzymes added by the honeybee during the ripening of honey or by chemical action in the concentrated, somewhat acid sugar mixture we know as honey.

Acids

The flavor of honey results from the blending of many “notes,” not the least being a slight tartness or acidity. The acids of honey account for less than 0.5 percent of the solids, but this level contributes not only to the flavor, but is in part responsible for the excellent stability of honey against microorganisms. Several acids have been found in honey, gluconic acid being the major one. It arises from dextrose through the action of an enzyme called glucose oxidase. Other acids in honey are formic, acetic, butyric, lactic, oxalic, succinic, tartaric, maleic, pyruvic, pyroglutamic, a-ketoglutaric, glycollic, citric, malic, 2- or 3-phosphoglyceric acid, a- or B-glycerophosphate, and glucose 6-phosphate.

Proteins and Amino Acids

It will be noted in table 1 that the amount of nitrogen in honey is low, 0.04 percent on the average, though it may range to 0.1 percent. Recent work has shown that only 40 to 65 percent of the total nitrogen in honey is in protein, and some nitrogen resides in substances other than proteins, namely the amino acids. Of the 8 to 11 proteins found in various honeys, 4 are common to all, and appear to originate in the bee, rather than the nectar. Little is known of many proteins in honey, except that the enzymes fall into this class.

The presence of proteins causes honey to have a lower surface tension than it would have otherwise, which produces a marked tendency to foam and form scum and encourages formation of fine air bubbles. Beekeepers familiar with buckwheat honey know how readily it tends to foam and produce surface scum, which is largely due to its relatively high protein content.

The amino acids are simple compounds obtained when proteins are broken down by chemical or digestive processes. They are the “building blocks” of the proteins. Several of them are essential to life and must be obtained in the diet. The quantity of free amino acids in honey is small and of no nutritional significance. Breakthroughs in the separation and analysis of minute quantities of material (chromatography) have revealed that various honeys contain 11 to 21 free amino acids. Proline, glutamic acid, alanine, phenylalanine, tyrosine, leucine, and isoleucine are the most common, with proline predominating.

Amino acids are known to react slowly, or more rapidly by heating, with sugars to produce yellow or brown materials. Part of the darkening of honey with age or heating may be due to this.

Minerals

When honey is dried and burned, a small residue of ash invariably remains, which is the mineral content. As shown in table 1, it varies from 0.02 to slightly over 1 percent for a floral honey, averaging about 0.17 percent for the 490 samples analyzed.

Honeydew honey is richer in minerals, so much so that its mineral content is said to be a prime cause of its unsuitability for winter stores. Schuette and his colleagues at the University of Wisconsin have examined the mineral content of light and dark honey. They reported the following average values:
Enzymes

One of the characteristics that sets honey apart from all other sweetening agents is the presence of enzymes. These conceivably arise from the bee, pollen, nectar, or even yeasts or micro-organisms in the honey. Those most prominent are added by the bee during the conversion of nectar to honey. Enzymes are complex protein materials that under mild conditions bring about chemical changes, which may be very difficult to accomplish in a chemical laboratory without their aid. The changes that enzymes bring about throughout nature are essential to life.

Some of the most important honey enzymes are invertase, diastase, and glucose oxidase.

Invertase, also known as sucrase or saccharase splits sucrose into its constitutent simple sugars, dextrose, and levulose. Other more complex sugars have been found recently to form in small amounts during this action and in part explain the complexity of the minor sugars of honey. Although the work of invertase is completed when honey is ripened, the enzyme remains in the honey and retains its activity for some time. Even so, the sucrose content of honey never reaches zero. Since the enzyme also synthesizes sucrose, perhaps the final low value for the sucrose content of honey represents an equilibrium between splitting and forming sucrose.

Diastase (amylase) digests starch to simpler compounds but no starch is found in nectar. What its function is in honey is not clear. Diastase appears to be present in varying amounts in nearly all honey and it can be measured. It has probably had the greatest attention in the past, because it has been used as a measure of honey quality in several European countries.

Glucose oxidase converts dextrose to a related material, a gulconolactone, which in turn forms gluconic acid, the principal acid in honey. Since this enzyme previously was shown to be in the pharyngeal gland of the honey bee, this is probably the source. Here, as with other enzymes, the amount varies in different honeys. In addition to gluconolactone, glucose oxidase forms hydrogen peroxide during its action on dextrose, which has been shown to be the basis of the heat-sensitive antibacterial activity of honey.

Other enzymes are reported to be present in honey, including catalase and an acid phosphatase. All the honey enzymes can be destroyed or weakened by heat.

Properties of Honey

Because of honey’s complex and unusual composition, it has several interesting attributes. In addition, honey has some properties, because of its composition, that make it difficult to handle and use. With modern technology, however, methods have been established to cope with many of these problems.

Antibacterial Activity

An ancient use for honey was in medicine as a dressing for wounds and inflammations. Today, medicinal uses of honey are largely confined to folk medicine. On the other hand, since milk can be a carrier of some diseases, it was once thought that honey might likewise be such a carrier. Some years ago this idea was examined by adding nine common pathogenic bacteria to honey. All the bacteria died within a few hours or days. Honey is not a suitable medium for bacteria for two reasons - it is fairly acid and it is too high in sugar content for growth to occur. This killing of bacteria by high sugar content is called osmotic effect. It seems to function by literally drying out the bacteria. Some bacteria, however, can survive in the resting spore form, though not grown in honey.

Another type of antibacterial property of honey is that due to inhibine. The presence of an antibacterial activity in honey was first reported about 1940 and confirmed in several laboratories. Since then, several papers were published on this subject. Generally, most investigators agree that inhibine (name used by Dold, its discoverer, for antibacterial activity) is sensitive to heat and light. The effect of heat on the inhibine content, of honey was studied by several investigators. Apparently, heating honey sufficiently to reduce markedly or to destroy its inhibine activity would deny it a market as first-quality honey in several European countries. The use of sucrase and inhibine assays together was proposed to determine the heating history of commercial honey.

Until 1963, when White showed that the inhibine effect was due to hydrogen peroxide produced and accumulated in diluted honey, its identity remained unknown. This material, well known for its antiseptic properties, is a byproduct of the formation of gluconic acid by an enzyme that occurs in honey, glucose oxidase. The peroxide can inhibit the growth of certain bacteria in the diluted honey. Since it is destroyed by other honey constituents, an equilibrium level of peroxides will occur in a diluted honey, its magnitude depending on many factors such as enzyme activity, oxygen availability, and amounts of peroxide-destroying materials in the honey. The amount of inhibine (peroxide accumulation) in honey depends on floral type, age, and heating.

A chemical assay method has been developed that rapidly measures peroxide accumulation in diluted honey. By this procedure, different honeys have been found to vary widely in the sensitivity of their inhibine to heat. In general, the sensitivity is about the same as or greater than that of invertase and diastase in honey.

Food Value

Honey is primarily a high-energy carbohydrate food. Because its distinct flavors cannot be found elsewhere, it is an enjoyable treat. The honey sugars are largely the easily digestible “simple sugars,” similar to those in many fruits. Honey can be regarded as a good food for both infants and adults.

The protein and enzymes of honey, though used as indicators of heating history and hence table quality in some countries, are not present in sufficient quantities to be considered nutritionally significant. Several of the essential vitamins are present in honey, but in insignificant levels. The mineral content of honey is variable, but darker honeys have significant quantities of minerals.

Granulation

Dextrose, a major sugar in honey, can spontaneously crystallize from any honeys in the form of its monohydrate. This sometimes occurs when the moisture level in honey is allowed to drop below a certain level.

A large part of the honey sold to consumers in the United States is in the liquid form, much less in a finely granulated form known as “honey spread” or finely granulated honey, and even less as comb honey. The consumer appears to be conditioned to buying liquid honey. At least sales of the more convenient spread form have never approached those of liquid honey.

Since the granulated state is natural for most of the honey produced in this country, processing is required to keep it liquid. Careful application of heat to dissolve “seed” crystals and avoidance of subsequent “seeding” will usually suffice to keep a honey liquid for 6 months. Damage to color and flavor can result from excessive or improperly applied heat. Honey that has granulated can be returned to liquid by careful heating. Heat should be applied indirectly by hot water or air, not by direct flame or high-temperature electrical heat. Stirring accelerates the dissolution of crystals. For small containers, temperatures of 140ºF for 30 minutes usually will suffice.

If unheated honey is allowed to granulate naturally, several difficulties may arise. The texture may be fine and smooth or granular and objectionable to the consumer. Furthermore, a granulated honey becomes more susceptible to spoilage by fermentation, caused by natural yeast found in all honeys and apiaries. Quality damage from poor texture and fermented flavors usually is far greater than any caused by the heat needed to eliminate these problems.

Finely granulated honey may be prepared from a honey of proper moisture content (17.5 percent in summer, 15 percent in winter) by several processes. All involve pasteurization to eliminate fermentation, followed by addition at room temperature of 5 to 10 percent of a finely granulated “starter” of acceptable texture, thorough mixing, and storage at 55º to 60ºF in the retail containers for about a week. The texture remains acceptable if storage is below about 80º to 85º.

Deterioration of Quality

Fermentation. - Fermentation of honey is caused by the action of sugar-tolerant yeasts upon the sugars dextrose and levulose, resulting in the formation of ethyl alcohol and carbon dioxide. The alcohol in the presence of oxygen then may be broken down into acetic acid and water. As a result, honey that has fermented may taste sour.

The yeasts responsible for fermentation occur naturally in honey, in that they can germinate and grow at much higher sugar concentrations than other yeasts, and, therefore, are called “osmophilic.” Even so there are upper limits of sugar concentration beyond which these yeasts will not grow. Thus, the water content of a honey is one of the factors concerned in spoilage by fermentation. The others are extent of contamination by yeast spores (yeast count) and temperature of storage.

Honey with less than 17.1 percent water will not ferment in a year, irrespective of the yeast count. Between 17.1 and 18 percent moisture, honey with 1,000 yeast spores or less per gram will be safe for a year. When moisture is between 18.1 and 19 percent, not more than 10 yeast spores per gram can be present for safe storage. Above 19 percent water, honey can be expected to ferment even with only one spore per gram of honey, a level so low as to be very rare.

When honey granulates, the resulting increased moisture content of the liquid part is favorable for fermentation. Honey with a high moisture content will not ferment below 50ºF or above about 80º. Honey even of relatively low water content will ferment at 60º. Storing at temperatures over 80º to avoid fermentation is not practical as it will damage honey.

E. C. Martin has studied the mechanism and course of yeast fermentation in honey in conjunction with his work on the hygroscopicity of honey. He confirmed that when honey absorbs moisture, which occurs when it is stored above 60-percent relative humidity, the moisture content at first increases mostly at the surface before the water diffuses into the bulk of the honey. When honey absorbs moisture, yeasts grow aerobically (using oxygen) at the surface and multiply rapidly, whereas below the surface the growth is slower.

Fermenting honey is usually at least partly granulated and is characterized by a foam or froth on the surface. lt will foam considerably when heated. An odor as of sweet wine or fermenting fruit may be detected. Gas production may be so vigorous as to cause honey to overflow or burst a container. The off-flavors and odors associated with fermentation probably arise from the acids produced by the yeasts.

Honey that has been fermented can sometimes be reclaimed by heating it to 150ºF for a short time. This stops the fermentation and expels some of the off-flavor. Fermentation in honey may be avoided by heating to kill yeasts. Minimal treatments to pasteurize honey are as follows:
The following summarize the important aspects of fermentation:

1. All honey should be considered to contain yeasts.

2. Honey is more liable to fermentation after granulation.

3. Honey of over 17 percent water may ferment and over 19 percent water will ferment.

4. Storage below 50ºF will prevent fermentation during such storage, but not later.

5. Heating honey to 150ºF for 30 minutes will destroy honey yeasts and thus prevent fermentation.

Quality loss by heating and storing - The other principal types of honey spoilage, damage by over-heating and by improper storing, are related to each other. In general, changes that take place quickly during heating also occur over a longer period during storage with the rate depending on the temperature. These include darkening, loss of fresh flavor, and formation of off-flavor (caramelization).

To keep honey in its original condition of high quality and delectable flavor and fragrance is possibly the greatest responsibility of the beekeeper and honey packer. At the same time it is an operation receiving perhaps less attention from the producer than any other and one requiring careful consideration by packers and wholesalers. To do an effective job, one must know the factors that govern honey quality, as well as the effects of various beekeeping and storage practices on honey quality. The factors are easily determined, but only recently are the facts becoming known regarding the effects of processing temperatures and storage on honey quality.

To be of highest quality, a honey - whether liquid, crystallized, or comb - must be well ripened with proper moisture content; it must be free of extraneous materials, such as excessive pollen, dust, insect parts, wax, and crystals if liquid; it must not ferment; and above all it must be of excellent flavor and aroma, characteristic of the particular honey type. It must, of course, be free of off-flavors or odors of any origin. In fact, the more closely it resembles the well-ripened honey as it exists in the cells of the comb, the better it is.

Several beekeeping practices can reduce the quality of the extracted product. These include combining inferior floral types, either by mixing at extracting time or removing the crop at incorrect times, extraction of unripe honey, extraction of brood combs, and delay in settling and straining. However, we are concerned here with the handling of honey from its extraction to its sale. During this time improper settling, straining, heating, and storage conditions can make a superb honey into just another commercial product.

The primary objective of all processing of honey is simple - to stabilize it. This means to keep it free of fermentation and to keep the desired physical state, be it liquid or finely granulated. Methods for accomplishing these objectives have been fairly well worked out and have been used for many years. Probably improvements can be made. The requirements for stability of honey are more stringent now than in the past, with honey a world commodity and available in supermarkets the year around. Government price support and loan operations require storage of honey, and market conditions also may require storage at any point in the handling chain, including the producer, packer, wholesaler, and exporter.

The primary operation in the processing of honey is the application and control of heat. If we consider storage to be the application of or exposure to low amounts of heat over long periods, it can be seen that a study of the effects of heat on honey quality can have a wide application.

Any assessment of honey quality must include flavor considerations. The objective measurement of changes in flavor, particularly where they are gradual, is most difficult. We have measured the accumulation of a decomposition product of the sugars (hydroxymethylfurfural or HMF) as an index of heat-induced chemical change in the honey. Changes in flavor, other than simple loss by evaporation, also may be considered heat-induced chemical changes.

To study the effects of treatment on honey, we must use some properties of honey as indices of change. Such properties should relate to the quality or commercial value of honey. The occurrence of granulation of liquid honey, liquefaction or softening of granulated honey, and fermentation as functions of storage conditions has been reported; also, color is easily measured.

As indicators of the acceptability of honey for table use, Europeans have for many years used the amount of certain enzymes and HMF in honey. They considered that heating honey sufficiently to destroy or greatly lower its enzyme content or produce HMF reduced its desirability for most uses. A considerable difference has been noted in the reports by various workers on the sensitivity to heat of enzymes, largely diastase and invertase, in honey. Only recently has it been noted that storage alone is sufficient to reduce enzyme content and produce HMF in honey. Since some honey types frequently exported to Europe are naturally low in diastase, the response of diastase and invertase to storage and processing is of great importance for exporters.

A study was made of the effects of heating and storage on honey quality and was based on the results with three types of honey stored at six temperatures for 2 years. The results were used to obtain predictions of the quality life of honey under any storage conditions. The following information is typical of the calculations based on this work.

At 68ºF, diastase in honey has a half-life of 1,500 days, nearly 4 years. Invertase is more heat sensitive, with a half-life at 68º of 800 days, or about 2-1/4 years. Thus there are no problems here. By increasing the storage temperature to 77º, half the diastase is gone in 540 days, or 1-1/3 years, and half the invertase disappears in 250 days, or about 8 months. These periods are still rather long and there would seem to be nothing to be concerned about. However, temperatures in the 90’s for extended periods are not at all uncommon: 126 days (4 months) will destroy half the diastase and about 50 days (2 months) will eliminate half the invertase. As the temperature increases, the periods involved become shorter and shorter until the processing temperatures are reached. At 130º, 2-1/2 days would account for half the diastase and in 13 hours half the invertase is gone.

A recommended temperature for pasteurization of honey is 145ºF for 30 minutes. At this temperature diastase has a half-life of 16 hours and invertase only 3 hours. At first glance this might seem to present no problems, but it must be remembered that unless flash heating and immediate cooling are used, many hours will be required for a batch of honey to cool from 145º to a safe temperature.

If we proceed further to a temperature often recommended for preventing granulation, 160ºF for 30 minutes, the necessity of prompt cooling becomes highly important. At 160º, 2-1/2 hours will destroy half of the diastase, but half of the more sensitive invertase will be lost in 40 minutes. This treatment then cannot be recommended for any honey in which a good enzyme level is needed, as for export.

The damage done to honey by heating and by storage is the same. For the lower storage temperatures, simply a much longer time is required to obtain the same result. It must be remembered that the effects of processing and storage are additive. It is for this reason that proper storage is so important. A few periods of hot weather can offset the benefits of months of cool storage - 10 days at 90ºF are equivalent to 100 to 120 days at 70º. An hour at 145º in processing will cause changes equivalent to 40 days’ storage at 77º.

An easy way for beekeepers to decide whether they have storage or processing deterioration is to take samples of the fresh honey, being careful that the samples are fairly representative of the batch, and place them in a freezer for the entire period. At the end of this time, they should warm the samples to room temperature and compare them by color, flavor, and aroma with the honey in common storage. In some parts of the United States, the value of the difference can reach 1-1/2 cents per pound in a few months. Such figures certainly would justify expenditures for temperature control.

People who store honey are in a dilemma. They must select conditions that will minimize fermentation, undesirable granulation, and heat damage. Fermentation is strongly retarded below 50ºF and above 100º. Granulation is accelerated between 55º and 60º and initiated by fluctuation at 50º to 55º. The best condition for storing unpasteurized honey seems to be below 50º, or winter temperatures over much of the United States. Warming above this range in the spring can initiate active fermentation in such honey, which is usually granulated and thus even more susceptible.

References

DONER, L. W.
1977. THE SUGARS OF HONEY-A REVIEW. Journal of Science and Agriculture.

TOWNSEND, G. F
1961. PREPARATION OF HONEY FOR MARKET. 24 p. Ontario Department of Agriculture Publication 544.

WHITE, J. W. JR.
1975. HONEY. In Grout, R. A., ed., The hive and the honey bee, p. 491-530. Dadant & Sons, Inc., Hamilton, Ill.

_________
1975. COMPOSITION AND PHYSICAL PROPERTIES OF HONEY. In E. Crane, ed., Honey Review, p. 157-239. Heinemann, London.

______ M. L. RIETHOP, M. H. SUBERS, and I. KUSHNIR.
1962. COMPOSITION OF AMERICAN HONEYS. 124 p. U.S. Department of Agriculture Technical Bulletin 1261.

By F. E. MOELLER(1)

BEEKEEPING IN THE UNITED STATES
AGRICULTURE HANDBOOK NUMBER 335
Revised October 1980
Pages 64 - 72

Colonies of bees existing in the wild, away from the control of human beings, will produce small surplus crops of honey above their requirements for survival. Such surplus will vary, depending on the region or locality, but will seldom exceed 25 to 30 pounds. In the same area and with the same nectar resources, colonies properly managed will produce surplus honey crops exceeding 100 pounds. Intensive two-queen colony management often can result in surplus crops of 300 pounds or more with the same resources available. The key to these differences is management.

Proper management employs practices that harmonize with the normal behavior of bees and brings the colony to its maximum population strength at the start of the bloom of major nectar-producing plants. Management practices are similar in basic principle wherever bees are kept and vary only as regards timing for the desired nectar source of the region or locality concerned.

Honey bee biology is constant. Bees respond to their environment as temperatures and food supplies are changed. Beekeepers, in managing or manipulating colonies, are merely facilitating normal biological colony changes to suit their purpose. They can accelerate brood rearing by pollen feeding and hive manipulation, or they can crowd or restrict colony activity by certain other manipulations. Responses of the colony, wherever it is kept, are predictable. Thus, the basic handling, management, and manipulation of bees are universally similar, varying only as to localities and the timing of bloom of the major nectar and pollen plants.

Regardless of the type of hives or equipment used, proper management aims at providing colonies with unrestricted room for brood rearing, ripening of nectar, and storage of honey, plus provision of adequate food requirements, both pollen and honey, for the time of year concerned. Swarming is minimized and the storing instinct encouraged when proper management is used.

(1) Research entomologist, Science and Education Administration (deceased).

Preparing Colony for New Season

In the temperate regions of the Northern Hemisphere, August to October is the time when beekeepers prepare their colonies for the coming year. This is when the major honey flows are usually past and the bees must be made ready for the coming winter.

All queens of questionable performance with only a small amount of brood of irregular pattern (fig. 1, A) should be replaced. Frequently, the bees of the colony will replace or supersede queens of subnormal performance even before the beekeeper senses a problem. Some queens may be satisfactory in their second year; queens less than a year old are usually best.

FIGURE 1. - Queens with (A) irregular and (B) good brood pattern.

To requeen a colony, certain principles of queen acceptance must be borne in mind: (1) Strong colonies more reluctantly accept a queen than weaker ones, (2) temperamental bees are more reluctant to accept a new queen than gentle bees, (3) young bees accept a queen more readily than older bees, (4) the colony to be requeened should first be made queenless, and (5) the queen to be introduced should be in egg-laying condition.

There is less risk in requeening a colony by giving it a laying queen with some of her own brood and bees than by giving it a queen in a shipping cage. A new or valuable queen should first be introduced into a small colony or divisions of one in a queen-shipping cage. After she is laying, the small colony can be united with a large one.

A drone-laying queen can be replaced if she is discovered while the colony is still strong. If the colony is weak, the bees should be removed and the equipment added to another colony.

Assuming colony conditions and the condition of the queen are favorable, the effect of environmental or working conditions and the time of year are factors that affect queen acceptance. Best acceptance is usually obtained when some nectar is available in the field.

One possible period for requeening is during the broodless period of late fall. Queens are easily introduced at this time, and the bees are passive to their presence. However, the uncertainty of the weather, the difficulty of finding old and shrunken queens, and the danger of inciting robbing make this time of year less desirable for requeening than the summer.

Brood rearing declines in late summer and fall, and many normal colonies are completely broodless during much of November and December, particularly if the colony has no pollen. Older queens stop brood rearing sooner than younger queens.

Brood rearing should be encouraged as late in the season as possible. This can be assured by providing vigorous young queens in late summer, by preventing undue overcrowding and restriction of the brood nest with honey, and by encouraging pollen storage.

In areas where fall honey flows occur, partially filled supers should be kept on the colonies, especially if the brood nest is heavy

If brood rearing is restricted by a crowded brood nest or because of poor queens, the colony may enter the winter with a high percentage of old bees that will die early in the winter. Such colonies may later develop serious nosema infections and perish before spring. A colony should start the winter with about 10 pounds of bees and plenty of honey to carry it to the next spring.

Beekeepers in certain localities will need to think of winter stores for their colonies as early as the first of August if later honey flows are not dependable or are nonexistent. In October, colonies should have at least 45 pounds of honey in dark combs in the top brood chamber and 20 to 30 pounds of honey in each of two lower hive bodies - a total of at least 90 pounds of honey.

Preparing Colony for Winter

Population

The strength of a colony of bees is relative and difficult to describe. A “strong” colony to one beekeeper might be “weak” to another. Colonies with less than 10 pounds of bees should be united to stronger ones or several weaker ones combined. At between 40º and 50ºF, 10 pounds of bees will cover practically all the combs of a three-story standard hive wall to wall and top to bottom. Naturally, as the temperature drops, the cluster will contract.

The beekeeper must see that at no time is the available space for brood rearing reduced because of overcrowding with honey from the fall flow. A balance must be maintained between crowding the colony to get the brood chambers well filled with honey and adding space to relieve brood-rearing restriction. Partly filled supers kept on colonies in the fall may be necessary. Any subnormal colony should not be overwintered but united with another colony.

A colony may appear to have an adequate fall population, but if the bees are old, it will weaken rapidly as winter advances and may starve to death. Starvation occurs even with abundant honey in the hive because the cluster is too small to cover the honey stores.

Food Reserves

The colony should have a minimum of 500 square inches of comb filled with pollen in the fall. To insure uninterrupted brood rearing in late winter and early spring, the beekeeper may need to supplement these stores. The average colony of bees under intensive management may consume about 60 pounds of honey between the last flow in the fall and the first available food from the field in the spring. A weak colony may consume 20 pounds or less, but the very best colony will consume 80 pounds or more. To insure the survival of the top-quality colony, 90 to 100 pounds of honey should be left on it in the fall. A colony of bees not rearing brood will average about one-eighth pound of honey a day or 4 pounds a month. When brood rearing begins, the consumption of honey is greatly accelerated. Brood rearing should start in midwinter and accelerate as temperatures moderate in late winter and early spring.

When brood rearing is discouraged or curtailed, the colony will consume less winter stores but will emerge in the spring much weaker and with a population of primarily old bees. Such colonies will have difficulty replacing the small amount of honey they used over winter, whereas other colonies that have had normal, unimpeded rearing of brood will soon be able to replace all the honey they consumed over winter plus a substantial surplus.

Organization

To accommodate the best queens in standard Langstroth 10-frame hives, a minimum of 2 hive bodies, preferably three, should be used for year-round management. In the fall, most of the honey should be located in the top hive body. With experience, the beekeeper can soon learn to estimate the weight of hive bodies or frames by lifting them. A frame full of honey should weigh approximately 5 pounds. The top hive body should contain 40 to 45 pounds of honey. This means that all frames in the top hive body will be full of honey except for two or three frames in the center. The second body should contain 25 to 30 pounds of honey and some pollen. The bottom hive body should contain 20 to 30 pounds of honey plus pollen. If in the fall the combs in the top hive body are not filled, the beekeeper should reorganize them and if necessary feed additional sugar syrup so that this top hive body is well filled with stores.

As the winter progresses, the cluster of bees will shift its position upward as the stores are consumed. A colony of bees in a cold climate can starve with abundant honey in the hive if the honey is below the cluster.

With the advent of cold weather, the bees cluster tightly in the interspaces of the combs. Usually there are no bees in the bottom part of the hive near the entrance. For this reason, an entrance cleat or reducer should he used to exclude mice, such as 1-inch auger holes drilled into the hive bodies of the brood nest just below the hand-holds. In late summer, these auger-hole entrances are closed with corks so that the bees will fill the combs near them. During winter the top auger-hole entrance should be open. This allows the escape of moisture-laden air and affords a flight exit for the bees during warm spells (fig. 2).

Packing the Hive

Many beekeepers in the coldest parts of the country consider that some form of protection around the hive is essential. Others believe that colonies with strong populations and ample stores need no further protection. Factors to consider in deciding whether to pack are the cost of material and labor and any savings in honey or bees. Packing will not replenish colonies deficient in honey, pollen, or bees, replace poor queens, or cure bee diseases. Packed colonies will consume slightly less honey. The difference, however, is negligible. The most important consideration in preparing colonies for winter is a strong population and adequate stores.

When outside temperatures are near freezing, the temperature at the surface of a cluster of bees ranges between 43º and 46ºF. As the temperature decreases, the cluster contracts and the bees in the outer insulating shell concentrate to provide an insulating band 1 to 3 inches in depth. Metabolism and activity of the bees in the center of the cluster maintain a desired temperature. This may be around 92º if brood rearing is in progress. The temperature of the area of the hive not occupied by bees will be similar to the external temperature. The difference is that the temperature in the unpacked hive changes more rapidly and responds more quickly to that outside the hive. Heavy packing is worse than no packing, because during warm periods in midwinter when the bees should fly, those heavily packed do not fly at all.

It is important to consider the strength of the colony so that the bees can, at all times, cover a good percentage of their winter stores. If the population becomes weakened so that they cannot cover more than a few pounds of honey at a time, they can starve to death because they do not have contact with sufficient food.

Late Winter Manipulation

If colonies are inspected in later winter or early spring, adjustments can be made to save colonies that might be lost otherwise. Even weak or medium-strength colonies often can be saved if honey is moved into contact with the cluster. A strong colony with insufficient honey can starve if additional food is not provided at this time.

From this period until the bees can forage, such colonies can be fed either full combs of honey, or if these are not available, a gallon or two of heavy sugar syrup (two parts sugar by volume to one part water) can be poured directly into the open cells of empty combs.

Spring Buildup

Overwintered colonies usually will start brood rearing in midwinter and continue into the summer unless the stored pollen is all consumed before fresh pollen is available. If the supply is exhausted and not supplemented, brood rearing will slow down or stop entirely when it should proceed without interruption.

For best results in honey production, a beekeeper should have strong populations of young bees for the honey flow. Colonies emerging in the spring with predominantly old bees must build a population of young bees for later flows by using the early sources of pollen.

Some beekeepers trap pollen at the hive entrance from incoming bees by means of a pollen trap such as that described in “Trapping Pollen From Honey Bee Colonies” (Detroy 1976). This pollen is dried or frozen until needed, then mixed with sugar, water, and soy flour, and fed to the colony as a supplement to its natural supply (fig. 3). Various other types of pollen supplements and substitutes have been described and some are available on the open market.

Supplements containing pollen are eaten more readily by bees and generally give better results than those containing soy flour or other material without pollen. Pollen supplement is preferred by the bees in direct proportion to the amount of pollen it contains. The less pollen the supplement contains, the less is eaten. Substitutes made without pollen tend to be dry and gummy. A pound of pollen will make approximately 12 pounds of pollen supplement.

Swarm Control in Single-Queen Management

After pollen becomes abundantly available in the spring, the beekeeper should provide ample space for brood rearing and honey storage.

The natural colony behavior is to expand its brood nest upward, and a simple manipulation utilizing this tendency is to shift the empty frames or emerging brood to the top of the hive and the youngest brood and honey to the bottom part. This permits the expansion of the brood rearing upward into this area (fig. 4). Subsequent reversal of brood chambers can be made at about 10-day or 2-week intervals until the honey flow starts.

As soon as the three brood chambers are filled with bees, the first super should be given whether or not the honey flow is in progress. If this is done, most colonies with a vigorous queen will not swarm. However, any queen cells the beekeeper sees as he reverses the brood chambers should be removed. A simple method of reversing brood chambers is to lower the hive backward to the ground, separate the brood chambers, interchange the first and third hive bodies, and return to position.

After the honey flow starts, the danger of swarming lessens and brood chamber reversal can be discontinued. At the start of the honey flow, “bottom supering” should be used. The empty super should be placed above the top brood chamber but below the partially filled supers (fig. 4).

After the supers are filled and the honey extracted, they should never be put directly over the brood nest, but should be placed on top of the partly filled supers to prevent the queen seeking them and laying eggs in them. Why such combs attract the queen is not known.

Two-Queen System

The establishment of a two-queen colony is based on the harmonious existence of two queens in a colony unit. Any system that ensures egg production of two queens in a single colony for about 2 months before the honey flow will boost honey production (Moeller 1956).

The population in a two-queen colony may be twice the population of a single-queen colony. Such a colony will produce more honey and produce it more efficiently than will two single-queen colonies. A two-queen colony usually enters winter with more pollen than a single-queen colony. As a result of this pollen reserve, the two-queen colony emerges in the spring with a larger population of young bees and is thus a more ideal unit for starting another two-queen system.

To operate two-queen colonies, start with strong overwintered colonies. Build them to maximum strength in early spring. Obtain young queens about 2 months before the major honey flows start. When the queens arrive, temporarily divide the colony. Replace the old queen, most of the younger brood, and about half the population in the bottom section. Cover with an inner cover or a thin board and close the escape hole. The division containing most of the sealed and emerging brood, the new queen, and the rest of the population is placed above. The upper unit is provided with an exit hole for flight.

At least two brood chambers must be used for the bottom queen and two for the top queen. Two weeks after the new queen’s introduction, remove the division board and replace it with a queen excluder. The supering is double that required for a single-queen operation, or where three standard supers are needed for a single colony, six will be needed for a two-queen colony.

When supering is required, larger populations in two-queen colonies require considerably more room at one time than is required for single-queen colonies. If a single-queen colony receives one super, a two-queen system may require two or even three empty supers at one time.

The brood chambers should be reversed to allow normal upward expansion of the brood area about every 7 to 10 days until about 4 weeks before the expected end of the flow, after which the honey crop on the colony may be so heavy as to preclude any brood nest manipulations. Thereafter, give supers as they are needed for storage of the crop. As the honey is extracted, the supers are returned to the hive to be refilled. They should never be replaced directly over the top brood nest, unless a second queen excluder is used to keep the queen out of them. The top brood nest may tend to become honey bound. If this occurs, reverse the upper and lower brood nests around the queen excluder. This puts the top honey-bound brood nest on the bottom board and the lighter brood nest with the old queen above the excluder.

There is no advantage in having a second queen when about a month of honey flow remains, because eggs laid from this time on will not develop into foragers before the flow has ended. However, entering the brood nest during the middle of the flow to remove one of the queens is impractical. Uniting back to a single-queen status can be done after the bulk of the honey is removed from the colony. By this time some colonies may have already disposed of one queen. When this happens, simply remove the queen excluder and operate the colony as a single-queen unit.

Improved Stock

Production of honey is one major criterion in selecting honey bee stock and breeding for improvement. Superior stock must also be reasonably gentle, not prone to excessive swarming, maintain a large but compact brood nest, and winter well. It should ripen its honey rapidly, seal the cells with white wax, and use a minimum of burr comb. To obtain all the desirable characters in a superior stock, specific inbred lines from many sources must be selected and developed and then recombined into a genetically controlled hybrid. When this is done, hybrid vigor or heterosis usually results (Moeller 1976).

Queens of common stock reared under favorable conditions and heading well-managed colonies probably will be more productive than poorly reared queens of superior stock. Queens of superior stock reared under favorable conditions will require a higher standard of management than is demanded of common stock. To realize the maximum benefits from improved stock, the beekeeper must provide unrestricted room for brood rearing, ripening of nectar, and storage of honey.

The queen breeder should produce the best queens possible to obtain the maximum benefits from improved stock and the honey producer receiving these queens should manage them in such a way that they can develop their maximum colony populations.

Disease Control as Affected by Good Management

If colonies are operated for highest honey yields, they must be kept in optimum condition (fig. 5). This includes rigid control of all bee diseases. For information about bee diseases, see pages 118 to 128.

FIGURE 5. - Brood combs showing (top) healthy brood necessary for high honey production and (bottom) diseased brood, which results in weakened colonies and low honey production.

References

DETROY, B. F. and E. R. HARP
1976. TRAPPING POLLEN FROM HONEY BEE COLONIES. 11 p. U.S. Department of Agriculture, Production Research Report 163.

FARRAR, C. L.
1937. INFLUENCE OF COLONY POPULATIONS ON HONEY PRODUCTION. Journal of Agricultural Research 54:945-954.

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1942. NOSEMA DISEASE CONTRIBUTES TO WINTER LOSSES AND QUEEN SUPERSEDURE. In Gleanings in Bee Culture 70:660-661, 701.

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1944. PRODUCTIVE MANAGEMENT OF HONEY BEE COLONIES IN THE NORTHERN STATES. 20 p. U.S. Department of Agriculture Circular 702.

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1973-74. PRODUCTIVE MANAGEMENT OF HONEY BEE COLONIES. American Bee Journal 113(8-12), Aug. through Dec. 1973; 114(1-3) Jan. through Mar. 1974.

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1952. ECOLOGICAL STUDIES ON OVERWINTERED HONEY BEE COLONIES. Journal of Economic Entomology 45:445-449.

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1960. OLD AND NEW IDEAS ABOUT WINTERING. American Bee Journal 100:306-310.

HOOPINGARNER, R., and C. L. FARRAR
1959. GENETIC CONTROL OF SIZE IN QUEEN HONEY BEES. Journal of Economic Entomology 52:547-548.

MOELLER, F. E.
1956. BEHAVIOR OF NOSEMA-INFECTED BEES AFFECTING THEIR POSITION IN THE WINTER CLUSTER. Journal of Economic Entomology 49:743-745.

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1961. THE RELATIONSHIP BETWEEN COLONY POPULATIONS AND HONEY PRODUCTION AS AFFECTED BY HONEY BEE STOCK LINES. 20 p. U.S. Department of Agriculture, Production Research Report 55.

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1976. TWO-QUEEN SYSTEM OF HONEY BEE COLONY MANAGEMENT. 11 p. U.S. Department of Agriculture, Production Research Report 161.

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1976. DEVELOPMENT OF HYBRID HONEY BEES. 11 p. U.S. Department of Agriculture, Production Research Report 168.

SCHAEFER, C. W. and C. L. FARRAR
1946. USE OF POLLEN TRAPS AND POLLEN SUPPLEMENTS IN DEVELOPING HONEY BEE COLONIES. 13 p. U.S. Bureau of Entomology and Plant
Quarantine, E-531 revised.

BY STEPHEN TABER III(1)

BEEKEEPING IN THE UNITED STATES
AGRICULTURE HANDBOOK NUMBER 335
Revised October 1980
Pages 33 - 38

Bee behavior refers to what bees do - as individuals and as a colony. By studying their behavior, we may learn how to change it to our benefit.

Two practical discoveries of bee behavior made our beekeeping of today possible. One was the discovery by Langstroth of bee space. The other was the discovery by G. M. Doolittle that large numbers of queens could be reared by transferring larvae to artificial queen cups. The discovery of the “language” of bees and of their use of polarized light for navigation has attracted considerable interest all over the world.

Much has been learned about the behavior of insects, including bees, in recent years. As an example, the term “pheromone” had not been coined in 1953, when Ribbands summarized the subject of bee behavior in his book, The Behaviour and Social Life of Honeybees. A pheromone is a substance secreted by an animal that causes a specific reaction by another individual of the same species. Now many bee behavior activities can be explained as the effect of various pheromones.

Recently, we have learned how certain bee behavior activities are inherited, and this information gives us a vast new tool to tailor-make the honey bee of our choice. Further studies should reveal other ways to change bees to produce specific strains for specific uses.

(1) Apiculturist, Science and Education Administration, Carl Hayden Center for Bee Research, Tuscon, Ariz. 85719.

The Honey Bee Colony

The physical makeup of a colony has been described. An additional requirement of a colony is a social pattern or organization, probably associated with a “social pheromone.” It causes the bees to collect and store food for later use by other individuals. It causes them to maintain temperature control for community survival when individually all would perish. Individuals within the colony communicate with each other but not with bees of another colony. Certain bees in the colony will sting to repel an intruder, even though the act causes their death. All of these, and perhaps many other organizational activities, probably are caused by pheromones.

There is no known governmental hierarchy giving orders for work to be done, but a definite effect on the colony is observed when the queen disappears. This effect seems to be associated with a complex material produced by the queen that we refer to as “queen substance.” There also is evidence that the worker bees from 10 to 15 days old, who have largely completed their nursing and household duties but have not begun to forage, control the “governmental” structure. Just what controls them has not been determined.

These and many other factors make an organized colony out of the many thousands of individuals.

The Domicile

When the swarm emerges from its domicile and settles in a cluster on a tree, certain “scout bees” communicate to it the availability of other domiciles. At least some of these domiciles may have been located by the scout bees before the swarm emerged. The various scouts perform their dances on the cluster to indicate the direction, distance, and desirability of the domiciles. Eventually, the cluster becomes united in its approval of a particular site. Then the swarm moves in a swirling mass of flying bees to it. Agreement always is unanimous.

When a swarm or combless package is placed in a box, allowed to fly, and supplied with abundant food, it builds comb. With a laying queen present, the first comb is “worker” in design, with about 25 cells per square inch. As the population of bees grows larger, and after there is a considerable amount of worker comb built, comb containing larger cells is constructed. This comb, termed storage comb by Langstroth, is used for rearing drones. We have found that bees store their first honey all across the top of the combs, utilizing both drone and worker cells.

The space between honey storage combs is much more uniform than between brood combs. The space left between capped honey cells is usually one-fourth inch or even less - room enough for one layer of bees to move.

As the colony ages, the combs that were first used for rearing worker bees may be converted to honey storage comb; areas damaged in any way are rebuilt. These changes usually affect the bee space and result in combs being joined together with “brace” comb. Strains of bees show genetic variation in building these brace combs.

All these cells are horizontal or nearly so; vertical cells are used for rearing queens. Why horizontal cells are used for the rearing of brood and for honey and pollen storage, whereas vertical cells are built only for queen production, is unknown.

Flight Behavior

When several thousand bees and a queen are placed in new surroundings - which happens when the swarm enters its new domicile or a package of bees is installed, or a colony is moved to a new location - normal flight of some workers from the entrance may occur within minutes. If flowering plants are available, bees may be returning to the hive with pollen within an hour. Bees transferred by air from Hawaii to Louisiana and released at 11:30 a.m. were returning to the new location with pollen loads within an hour. Package bee buyers in the Northern States have noticed similar patterns in bees shipped from the South.

What causes this virtually instant foraging by bees? What determines whether they collect pollen, nectar, or water? If food and water in the hive are sufficient, why should they leave to forage? Answers to these questions may lead to our directing bees to specific duties we desire accomplished.

Housecleaning

Certain waste material accumulates in a normal colony. Adult bees and immature forms may die. Wax scales, cappings from the cells of emerging bees, particles of pollen, and crystallized bits of honey drop to the floor of the hive. Intruders, such as wax moths, bees from other colonies, and predators, are killed and fall to the floor. Worker bees remove this debris from the hive.

The cleaning behavior of some strains of bees, associated with removal of larvae and pupae that have died of American foulbrood, is genetically controlled by two genes. This discovery is important not only because it might help in developing bees resistant to diseases, but also in indicating that other behavior characteristics of bees can be genetically modified to suit special needs.

Known Pheromone Activity

Chemicals that bees and other insects produce that influence, or direct, behavior of other bees are broadly called pheromones. In honey bees these chemicals are produced by the queen, workers, and probably drones. A list of the known chemicals associated with the queen and worker is given in tables 1 and 2.

This is an interesting and new area for bee research, as this list represents just a beginning. Research has indicated the existence of many other pheromones, which are as yet undocumented. If interested in this topic, consult the technical work listed in Gary (1974).

TABLE 1. - Queen pheromones
TABLE 2. - Worker bee pheromones

Pheromonal bee behavior activity patterns are easily observable. Nassanoff or scent gland activity is best seen when a swarm is hived. When the bees first enter the new domicile, some bees stand near the entrance and fan. At the same time, they turn the abdominal tip downward to expose a small, wet, white material on top of the end of the abdomen. This seems to affect the other bees, for within several minutes all will have entered the new hive. When bees find a new source of food, they also mark it with the same chemical.

Colony odor refers to the odor of one colony. Because each colony odor is different, colonies cannot be combined into one hive without the bees fighting and killing one another. This odor probably results from a combination of endogenous (pheromone or pheromonelike) materials and exogenous (food) materials in each hive and seems to be recognizably different for every colony.

When colonies are to be combined, the beekeeper usually places a newspaper between the two sets of bees. By the time the bees have eaten through and disposed of the newspaper, their odors have intermingled and become indistinguishable. During heavy honey flows, differences between colonies seem to disappear, or be submerged by the scent of nectar, and colonies can be united without difficulty.

One of the most interesting and complex pheromones, originally termed “queen substance,” is now believed to be a complex of different chemical pheromone compounds which stimulate a large number of complex behavior responses. Its presence in virgin queens in flight attracts the drone for mating from an unknown distance. Its presence in virgin and mated queens prevents the ovaries of the worker bee within the hive from developing and the worker bees from building queen cells. It keeps swarming bees near the queen. Its decrease is a cause of swarm preparation or supersedure. Queen substance is produced in glands in the queen’s head. The alarm or sting pheromone also may be a complex of pheromones. When a bee stings, other bees in the immediate vicinity also try to sting in the same place. Smoke blown onto the area seems to neutralize this effect.

Cause of Stinging Bees or Temper

The term “temper” of bees refers to their inclination to sting. Many factors influence the temper of bees, and it is a difficult subject to study. Environment of the hive and manipulation by an individual beekeeper certainly influence temper responses of bees. Temper is probably influenced tremendously by the genetics or inheritance of the bee as well as the environment. The Brazilian or Africanized bee is thought to be more genetically prone to sting than bees in the United States.

Temper of bees commonly has been controlled with smoke. Just why and how smoke affects bees is unknown, even though it has been used by beekeepers worldwide for hundreds of years. Furthermore, instructing beginners and novices exactly when and how to use smoke on bees is almost impossible. It is something that is learned from experience.

The following brief instruction might help beekeepers with limited experience: Smoke the entrance gently enough to force guard bees inside, raise cover, smoke gently. Smoke bees only when they fly up from combs toward hands and face. Move slowly and deliberately. Break propolis seals between hive bodies and frames slowly and evenly.

Don’t jar or bump combs and bees. During cold weather, propolis joints snap when pried apart un-ess care is taken. If combs are kept clean of propolis and burr and brace comb and if care is taken not to crush bees when moving combs and supers, they can be kept quite gentle.

Great care should be exercised in the placement of colonies of bees so that they cannot become a nuisance to friends and neighbors. Bees visiting nearby fishponds, swimming pools, and stock-watering troughs can be a real nuisance as well as dangerous to people and animals. Springtime flight of bees voiding feces and spotting laundry hanging on a line or a new car is irritating. Good public relations are important for beekeepers. Talk to your neighbors about the importance of bees in the community and country at large. Help them to understand that your bees and others are responsible for important pollination and share some honey with them occasionally.

Colony Morale

“Colony morale” generally refers to the well-being of the colony. If the morale is good, the bees are doing what is desired of them, including increasing the colony population, making honey, and pollinating flowers. Many factors affect colony morale. For example, if the queen is removed from a colony during a honey flow, the daily weight gains immediately decrease, although the bee population for the next 3 weeks is unaltered. Also, when a colony is preparing to swarm, the bees practically stop gathering pollen and nectar. Improper manipulations or external environment also affects colony morale. A colony has good morale when the maximum number of bees are making the maximum number of flights to gather nectar and pollen.

Other Methods of Bee Communication

There are other methods of bee communication besides the one involving chemical pheromones. The best known is the “dance” of the returned forager bee so well elucidated by von Frisch and his many students, particularly M. Lindaner.

This dance is so precise that it tells other bees not only in which direction to go but also how far to fly in search of food. This was the first non-human language to be interpreted. The experiments on bee communication by dances were done with dishes of sugar water and not under true foraging conditions of bees collecting nectar from plants. When a returning forager comes back to the hive after finding a highly attractive 100-acre field of sweetclover, does she direct bees to the spot she was working or to the whole field? The last word in dance communication of bees certainly has not yet been written.

Even the most uninitiated are familiar with the soft quiet hum of bees collecting nectar and pollen on their foraging trips. In the hive itself, there are many more bee noises or sounds which are much more subtle. Experienced beekeepers recognize a difference in sound between a colony with a queen and one without. Individual queens and even worker bees emit squeaky sounds called “piping” and “quacking.” The bee literature is full of many explanations of the causes and meanings of these sounds. Since these sounds and other hive sounds are now under careful scientific scrutiny, it is really premature to say definitely that they have certain defined meanings. This field of interest may produce useful information in the future.

According to von Frisch, when a bee returns from a foraging trip and dances, she also communicates the kind of “plant” or “flower” on which she was foraging by releasing the perfume of the flower through nectar regurgitation or from nectar aroma on body hairs. Again, most of these experiments were done with dishes of sugar water impregnated with essential oils or plant extracts. These experiments have prompted other experiments that were designed to train bees to work desired crops for pollination. These experiments were unsuccessful. The reason for the failures may well be that the bee language code has not been completely translated. We are still unable to “talk” effectively to the bees and “tell” them what we want done.

Von Frisch also discovered that bees recognize and are guided to flowers by different colors but are unable to communicate these colors. He also showed that the bee’s eyes are receptive to polarized light and that polarization of the light from the sky aids the bee’s navigation. How light of different wave lengths or intensity affects what goes on inside a hive is being studied.

Age Levels of Bees Correlated With Work Habits

The honey bee is adaptable to many environments. Honey bees that were native only to Europe, Asia, and Africa have adapted well to all but the polar regions of the world. Part of this adaptability lies in the capacity of the individual bee to “sense” what must be done, then to perform the necessary duty.

Under normal conditions, all ages of bees are in the hive and, in general, the bee’s age determines its daily activity. In response to special needs of the colony, however, bees are capable of altering the division of labor according to age. Young bees feed larvae, build comb, and ripen nectar into honey in a rather definite sequence. After about 3 weeks, they become field bees. If many field bees are killed by pesticides, young bees go to the field at a younger age to get necessary chores accomplished.

The Performance of Colonies

Genetically, we found that some bees produce more honey than others, but we do not know why. The individual bee may collect more because of its own genetic inheritance. The colony may store more honey because of the queen’s inherited ability to lay more eggs, resulting in a greater total population of bees in the hive, or because the bees are inherently longer lived.

We can affect the bee’s environment in conjunction with its inheritance, and our aim is to have good-quality bees and maintain the best colony morale possible. A beekeeper’s disturbance of the colony during the honey flow results in a marked decrease in the amount of honey stored for that day and even the following day. Colonies of bees should not be needlessly disturbed; however, some manipulation associated with many aspects of management is necessary.

Bee behavior toward different plants varies greatly. Some plants are particularly attractive for nectar or pollen; others are not. Strains of bees can be genetically selected to visit certain plants, and plants can be selected to be more attractive to bees. Attractive nectar or pollen, or both, can be important in ensuring pollination of bee-pollinated crops. Nectar and pollen availability in plants can be accidentally eliminated by breeding. When this occurs, there is a loss of a potential honey crop, but more important can be the loss of a seed or fruit crop because the plant no longer attracts pollinators. If plants such as soybeans, which cover enormous acreages, could be made more attractive to bees, honey and possibly soybean yields could be greatly increased.

A behavior characteristic of honey bees limits their effectiveness in pollinating some crops. Individual bees usually confine their foraging area in a series of trips to the field to a relatively small area such as a single fruit tree. On the other hand, the foraging area of a colony may comprise several square miles; honey bees flying 2.5 miles in all directions from a single hive have access to 12,500 acres. This characteristic and the fact that honey bees distribute themselves well over the area within flight range are important in locating and harvesting available nectar and pollen.

Control of Foraging

A major crop pollination goal is to control foraging bees and get them to more effectively visit and pollinate crops; conversely, we would like to repel them from areas where there is danger from insecticides or where they endanger people. Work with other insects - both social and nonsocial - indicates that this might be accomplished some day by chemical and physical means.

There is considerable evidence that different plant species produce varying attractant compounds associated with their nectar and pollen. Bees are highly attracted to the scent of recently extracted honeycomb and to the scent of honey being extracted or heated. Obviously, chemical scents of certain flowers and to some extent scents incorporated in the collected honey are attractive to bees or associated with available food.

Some pollens also contain chemical compounds that stimulate collection response in bees. Isolation and identification of these bee-attractive compounds and the application of the attractant to plant areas or altering attractants through plant breeding are an area of research of potential importance to crop pollination.

Research should not be confined to chemicals alone, but should be shared equally with various physical factors that can possibly attract or repel bees. In other entomological fields, research on physical methods of controlling insects is receiving intensive investigation. Different insects respond in differing ways; they are attracted to certain light wavelengths and repelled by others. Night-flying moths are repelled or go into defensive maneuvers because of bat sonar signals, whereas crickets and other members of their insect group can be collected by reproducing certain stridulations.

Other Behavior Activities of Bees

The Drones
The time of day that drones fly in search of a mate depends on many factors, such as the geographic location, day length, and temperature. Drones usually fly from the hive in large numbers between 11 a.m. and 4:30 p.m. Morning or early afternoon flights may last 2 or 3 hours. Later flights are shorter. When out of the hive, drones congregate in “mating areas,” which may serve to attract virgin queens. These areas usually are less than 100 feet from the ground and seem to be associated with land terrain.

The Queen
The virgin queen becomes sexually mature about 5 days after emergence. She is relatively quiet in the morning and most active in the afternoon. She may begin her mating flights 5 or 6 days after emergence and go on a number of flights over several days. Mating with 8 to 12 drones will stock her spermatheca with 6 million to 7 million sperm. She will begin to lay eggs in 2 to 5 days and may continue for years.

A young, fully mated queen rarely lays drone eggs before she is several months old. After that time, she controls the sex of the offspring by laying either fertilized or nonfertilized eggs.

Worker bees occasionally kill their queen. More frequently, they will kill a newly introduced or virgin queen. To do this, 15 or 20 worker bees collect about her in a tight ball until she starves. Generally, it has been thought that bees “balled” strange or introduced queens because they did not have the proper “colony” odor. The reason for balling is probably more complicated than that, because bees occasionally will ball their own queen. Even if the ball is broken up, the queen seldom survives and the stimulus is powerful enough that the bees taking part in the queen balling are sometimes subsequently balled by other bees.

References

BUTLER, C. G.
1955. THE WORLD OF THE HONEY BEE. 226 p. Macmillan Co., New York.

VON FRISHH, K.
1955. THE DANCING BEES. 183 p. Harcourt, Brace & Co., New York.

GARY, N. E.
1974. PHEROMONES THAT AFFECT THE BEHAVIOR AND PHYSIOLOGY OF HONEY BEES. In Pheromones, M. C. Birch, p. 200-221, North-Holland, Amsterdam, and Elsevier, New York.

HAYDAK, M. H.
1963. ACTIVITIES OF HONEY BEES. In The Hive and the Honey Bee, 556 p. Dadant & Sons, Hamilton, Ill.

LINDAUR, M.
1961. COMMUNICATION AMONG SOCIAL BEES. 143 p. Harvard University Press, Cambridge.

RIBBANDS, C. R.
1953. THE BEHAVIOUR AND SOCIAL LIFE OF HONEY BEES. 318 p. Dover Publications, Inc., New York.

By NORBERT M. KAUFFELD(1)

BEEKEEPING IN THE UNITED STATES
AGRICULTURE HANDBOOK NUMBER 335
Revised October 1980
Pages 30 - 32

A colony of honey bees comprises a cluster of several to 60,000 workers (sexually immature females), a queen (a sexually developed female), and, depending on the colony population and season of year, a few to several hundred drones (sexually developed males) (fig. 1). A colony normally has only one queen, whose sole function is egg laying. The bees cluster loosely over several wax combs, the cells of which are used to store honey (carbohydrate food) and pollen (protein food) and to rear young bees to replace old adults.

The activities of a colony vary with the seasons. The period from September to December might be considered the beginning of a new year for a colony of honey bees. The condition of the colony at this time of year greatly affects its prosperity for the next year.

In the fall a reduction in the amounts of nectar and pollen coming into the hive causes reduced brood rearing and diminishing population. Depending on the age and egg-laying condition of the queen, the proportion of old bees in the colony decreases. The young bees survive the winter, while the old ones gradually die. Propolis collected from the buds of trees is used to seal all cracks in the hive and reduce the size of the entrance to keep out cold air.

When nectar in the field becomes scarce, the workers drag the drones out of the hive and do not let them return, causing them to starve to death. Eliminating drones reduces the consumption of winter honey stores. When the temperature drops to 57º F, the bees begin to form a tight cluster. Within this cluster the brood (consisting of eggs, larvae, and pupae) is kept warm - about 93º F - with heat generated by the bees. The egg laying of the queen bee tapers off and may stop completely during October or November, even if pollen is stored in the combs. During cold winters, the colony is put to its severest test of endurance. Under subtropical, tropical, and mild winter conditions, egg laying and brood rearing usually never stop.

FIGURE 1. - Worker, queen, and drone bees.

As temperatures drop, the bees draw closer together to conserve heat. The outer layer of bees is tightly compressed, insulating the bees within the cluster. As the temperature rises and falls, the cluster expands and contracts. The bees within the cluster have access to the food stores. During warm periods, the cluster shifts its position to cover new areas of comb containing honey. An extremely prolonged cold spell can prohibit cluster movement, and the bees may starve to death only inches away from honey.

The queen stays within the cluster and moves with it as it shifts position. Colonies that are well supplied with honey and pollen in the fall will begin to stimulatively feed the queen, and she begins egg laying during late December or early January - even in northern areas of the United States. This new brood aids in replacing the bees that have died during the winter. The extent of early brood rearing is determined by pollen stores gathered during the previous fall. In colonies with a lack of pollen, brood rearing is delayed until fresh pollen is collected from spring flowers, and these colonies usually emerge from winter with reduced populations. The colony population during the winter usually decreases because old bees continue to die; however, colonies with plenty of young bees produced during the fall and an ample supply of pollen and honey for winter usually have a strong population in the spring.

(1) Research entomologist, Science and Education Administration, Carl Hayden Center for Bee Research, Tucson, Ariz. 85719.

Spring Activity

During early spring, the lengthening days and new sources of pollen and nectar stimulate brood rearing. The bees also gather water to regulate temperature and to liquefy thick or granulated honey in the preparation of brood food. Drones will be absent or scarce at this time of the year.

Later in the spring, the population of the colony expands rapidly and the proportion of young bees increases. As the population increases, the field-worker force also increases. Field bees may collect nectar and pollen in greater amounts than are needed to maintain brood rearing, and surpluses of honey or pollen may accumulate (fig. 2).

As the days lengthen and the temperature continues to increase, the cluster expands further and drones are produced. With an increase in brood rearing and the accompanying increase in adult bees, the nest area of the colony becomes crowded. More bees are evident at the entrance of the nest. A telltale sign of overcrowding is to see the bees crawl out and hang in a cluster around the entrance on a warm afternoon.

Combined with crowded conditions, the queen also increases drone egg laying in preparing for the natural division of the colony by swarming. In addition to rearing workers and drones, the bees also prepare to rear a new queen. A few larvae that would normally develop into worker bees are fed a special gland food called royal jelly, their cells are reconstructed to accommodate the larger queen, and her rate of development is speeded up. The number of queen cells produced varies with races and strains of bees as well as individual colonies.

Regardless of its crowded condition, the colony will try to expand by building new combs if food and room are available. These new combs are generally used for the storage of honey, whereas the older combs are used for pollen storage and brood rearing.

FIGURE 2. - A field bee returning to the hive with a load of pollen.

Swarming

When the first virgin queen is almost ready to emerge, and before the main nectar flow, the colony will swarm during the warmer hours of the day. The old queen and about half of the bees will rush en masse out the entrance. After flying around in the air for several minutes, they will cluster on the limb of a tree or similar object (fig. 3). This cluster usually remains for an hour or so, depending on the time taken to find a new home by scouting bees. When a location is found, the cluster breaks up and flies to it. On reaching the new location, combs are quickly constructed, brood rearing starts, and nectar and pollen are gathered. Swarming generally occurs in the Central, Southern, and Western States from March to June, although it can occur at almost any time from April to October.

After the swarm departs, the remaining bees in the parent colony continue their field work of collecting nectar, pollen, propolis, and water. They also care for the eggs, larvae, and food, guard the entrance, and build combs. Emerging drones are nurtured so that there will be a male population for mating the virgin queen. When she emerges from her cell, she eats honey, grooms herself for a short time, and then proceeds to look for rival queens within the colony. Mortal combat eliminates all queens except one. When the survivor is about a week old, she flies out to mate with one or more drones in the air. The drones die after mating, but the mated queen returns to the nest as the new queen mother. Nurse bees care for her, whereas prior to mating she was ignored. Within 3 or 4 days the mated queen begins egg laying.

FIGURE 3. - Hiving a swarm.

During hot summer days, the colony temperature must be held down to about 93º F. The bees do this by gathering water and spreading it on the interior of the nest, thereby causing it to evaporate within the cluster by its exposure to air circulation.

During the early summer, the colony reaches its peak population and concentrates on the collection of nectar and pollen and the storage of honey for the coming winter. After reproduction, all colony activity is geared toward winter survival. Summer is the time for storage of surplus food supplies. The daylight period is then longest, permitting maximum foraging, although rain or drought may reduce flight and the supply of nectar and pollen available in flowers. It is during the summer that stores are accumulated for winter. If enough honey is stored, then the beekeeper can remove a portion and still leave ample for colony survival.

BY WILLIAM P. NYE(1)

BEEKEEPING IN THE UNITED STATES
AGRICULTURE HANDBOOK NUMBER 335
Revised October 1980
Pages 10 - 15

Based on flora, beekeeping methods, and land topography, the continental United States can be divided into seven geographical regions (fig. 1). Each region is discussed here from the standpoint of honey production and methods of beekeeping operations.

The flora, climate, and nature of the terrain determine the system of management practiced by the beekeeper. For example, in the Appalachicola swamps of the Southeast, hives are placed on scaffolding to protect them from flood waters. In the Southwest, shade must be provided to protect the hives from the hot sun. Colonies in the north and mountainous areas must he protected from the cold, in certain forested areas from bears, and on the desert from drifting sand.

Beekeepers must pay for some locations; others are furnished free. Where bees are desired for pollination, the beekeepers usually are paid for their services.

Most beekeepers move colonies at night when the bees are all inside the hive. But when daytime temperatures exceed 43.3ºC (110ºF) in the Southwest, bees stay inside the hive and are more easily moved at midday than at night when they tend to cluster at the entrance.

FIGURE 1. - Beekeeping regions of the United States.

(1) Retired, formerly apiculturist, U.S. Department of Agriculture.

Northeast

The severe winters, short summers, and hilly or mountainous nature of the Northeast produce a variety of plants - but none which serves as a major source of nectar. However, alfalfa is becoming an important source of nectar in certain areas as new and better varieties are developed. Nectar from white clover, basswood, black locust, birdsfoot trefoil, various berries, and wild flowers contribute to producing a mixture of honey, much of which is sold locally to residents acquainted with the type produced, and some of the highest prices for honey are obtained here. Few commercial beekeepers operate in the Northeast.

Average honey production per colony is only 13.3 kg (29 lb), but occasionally locations where alfalfa is grown produce much higher averages. An estimated 175,000 colonies are in this region.

The colonies are seldom moved, except the few belonging to commercial or semicommercial beekeepers who may rent their bees for pollination of blueberries, cranberries, other fruits, or cucumbers. Many commercial beekeepers now remove most of the honey, and each hive is reduced to a two-story brood nest that is trucked to the Southeast where it is allowed to build up and be divided to form new colonies. The hives are returned to the Northeast in the spring for fruit pollination before the main honey flow.

Colonies that are not moved South are located where there are good air drainage, protection from cold winds, and exposure to as much winter sun as possible. For additional protection from cold winters, many colonies are “packed,” that is, wrapped with insulation and tar paper, leaving only the entrance exposed. Winter loss is usually high and is replaced with package bees and queens purchased from southern beekeepers. Shade in summer is unnecessary.

Most beekeepers overwinter their colonies in two- or three-story, 10-frame standard Langstroth hives. Two basic types of hive covers and bottom boards in use are the telescope cover and reversible bottom board, and the California-style top and bottom. The telescope covers create problems when hives are moved because the hives do not fit closely together on a truck and break open when roped tightly in place. Where migratory beekeeping is practiced, the California-style top and bottom are used as they permit better stacking of hives on a truck. When the honey flow starts, beekeepers add one or two deep supers for surplus honey storage or one or two shallow supers for section or comb honey production.

North-Central Region

The bulk of the honey from the north-central region comes from alfalfa, soybeans, sweetclovers (yellow and white), and the true clovers (alsike, ladino, red, and white), with minor surpluses from basswood, black locust, and raspberry. All of this is high-quality honey. Alfalfa and clover are the predominant American honeys. Less desirable grades come from aster, goldenrod, and smartweed. The variety of other plants, however, ensures something for the bees to work on from spring until frost. The bulk of comb honey produced by bees in 1-pound sections comes from this region.

There are approximately 918,000 colonies, many of which belong to commercial beekeepers. Average production of surplus honey per colony is 24 kg (52 lb).

Some colonies are killed in the fall, and the equipment is stored; then the hives are restocked in the spring with packages of bees and a queen purchased from southern beekeepers. Other colonies are wrapped with insulation and tar paper for winter protection. Some are left with ample stores of honey and pollen in locations protected from wind and exposed to warming sunlight (fig. 2). Still others have most of the honey removed, and the hives are reduced to two-story brood nests that are trucked to the South, where they are allowed to build up and be divided to form new colonies. These are then returned to the North in the spring. Midsummer shade is beneficial. Migratory beekeeping is increasing as beekeepers move their colonies from one location to another to take advantage of the various nectar flows.

Some colonies are rented for pollination of fruits, legumes, and cucumbers.

Southeast

Average production of honey per colony in this region, 14 kg (30 lb), is about the same as in the Northeast but less than elsewhere. An estimated 1,483,000 colonies are located permanently in the Southeast. In addition, many thousands of colonies are trucked in from the northern areas during the winter, then returned to the North in the spring.

Most U.S. queen breeders and package bee shippers are located in the Southeast. An estimated 300,000 kg (660,000 lb) of live bees and many thousands of queens are shipped from the Southeast annually. Some northern beekeepers pick up their package bees and queens in van-type, air-conditioned trucks for safe transportation to their northern locations.

FIGURE 2. - Apiary sheltered by hardwood forest in north-central region.

Except for sizable areas in Florida, little pollination is provided on a cost basis in this region. Bees are rented for occasional pollination of fruit orchards and legume seed and melon production. In Florida, bees are rented for citrus, cucurbits, melons, and other fruits and vegetables.

In the mountainous area, sourwood is the prevailing source of quality honey, along with tulip-poplar and clovers. Sourwood honey is almost water white, does not granulate readily, and is so esteemed that it usually passes directly from producer to consumer at far above the price of other honeys. Various other honeys, from light to dark and from mild to strong, are produced in the Southeast.

In the lower elevations, gallberry becomes the predominant nectar source. In the Appalachicola swamp area, tupelo, famous for its high levulose content and nongranulating characteristics, also is an excellent source of honey. Farther south in Florida, citrus is the major source, with clovers the major source toward the Mississippi Delta, where cotton also becomes important.

Considerable migratory beekeeping occurs, for the long season permits harvest of a crop of honey in one area before another harvest starts elsewhere.

Chunk honey production is common - that is, a chunk of comb honey in a jar of liquid honey. Little section honey is produced.

Preparing bees for winter requires little work. Bees usually are wintered in two- or three-story hives. The problem is to have ample stores of honey and pollen in the colony in the fall. This is necessary for the strong colonies needed in the early spring for package bee production or the early honey flows.

Colonies benefit from shade during the summer in the Southeast, and shade is essential in the southern part for maximum colony production.

Plains Region

The bulk of the honey from the plains region comes from sweetclover and alfalfa; much of it is produced by commercial beekeepers.

In this region, about 476,000 colonies produce 25 kg (55 lb) of honey per colony. Colonies are wintered and operated similarly to colonies in the north-central region. Shade is not generally necessary, although partial or thorough shading during extremely hot midsummer days is beneficial. Some of the highest production per colony is obtained in the plains region. One reason is that the sweet-clover and alfalfa fields are relatively large and can support many colonies, and many of the apiaries belong to commercial beekeepers.

Some of the colonies are trucked to southern areas for the winter; some are packed; some are killed and then restocked in the spring; and others receive no special winter treatment.

Colonies are used to a limited extent in the pollination of alfalfa, sweetclover, and cucumbers.

From this region westward to the Pacific, where migratory beekeeping is practiced to a greater extent than elsewhere, the California-style top and bottom rather than the telescoping top and reversible bottom are used, as they permit better stacking of colonies on trucks.

Mountain Region

The major source of honey in the mountainous region is alfalfa (figs. 5 and 6). About 330,000 colonies produce on an average 30 kg (66 lb) of honey per colony. More than two-thirds of the colonies belong to commercial beekeepers; some manage 2,000 colonies or more with only part-time summer help.

Honey production is almost entirely dependent on irrigation, although alfalfa now is grown on dry land. Some of the highest production per colony is obtained in the mountainous region. One reason for this is that the alfalfa fields are relatively large and can support many colonies (fig. 3). Weed spraying has reduced the sweetclover acreage, but sweetclover is now on the increase in some areas.

In migratory beekeeping from this area west and south, the colonies usually are moved at night. The hive entrances are not closed, but the truckload usually is covered with a plastic screen for long trips. Some colonies are packed during the winter, which is extremely cold and dry. Colonies not packed are located where they have good wind protection, exposure to the sun, and good air drainage. Spring buildup is slow and fall nectar flows are rare. Shading is unnecessary.

FIGURE 3. - Unsheltered colonies located for alfalfa honey production and pollination in Utah.

Migratory beekeeping is extensive. For example, probably no other region in the country can compare with the Delta area of central Utah with so many colonies (20,000 to 40,000) moved in from such long distances in so short a period. The region produces a major portion of the alfalfa seed in Utah (fig. 4). Many colonies are moved south or west for the badly needed spring buildup, then returned for the summer flow. Some colonies are killed in the fall and restocked in the spring.

Southwest

In this hot, semiarid region, there are 155,000 colonies that produce 21.4 kg (47 lb) of honey per colony. The major sources of nectar are alfalfa, cotton, and mesquite. Other sources include citrus, catclaw, tamarix, safflower, wild buckwheat, and other desert shrubs.

Summer shade is highly important (fig. 5). Artificial shade is often provided. Winter protection is unnecessary. Some colonies are wintered in a single brood nest with one or two shallow supers, but most are in two or three standard hive bodies. Nearby water is essential, and if it disappears even for only a day, the colonies may perish. Migration from one honey flow to another is common.

FIGURE 4. - Colonies in groups of 8 to 12 are placed one-tenth mile apart in large alfalfa seed fields for pollination.

FIGURE 5. - Typical apiary under a ramada that partially shades colonies in hot Southwest.

Colonies are used extensively in pollination of alfalfa and melons and to a lesser degree for citrus, onions, and cotton. A few package bees and queens are produced, but for the most part bees are kept for production of honey by commercial operators. Apiaries of 100 colonies or more are not unusual.

West

About 668,000 colonies in this region produce 12 kg (27 lb) of honey per colony. This production is rather meaningless because of the differences due to extreme variations in temperature, rainfall, elevation, and flora. The main source of nectar is alfalfa, which produces a light-colored honey of excellent flavor. Cultivated field crops such as clover, citrus, cotton, lima beans, deciduous fruit trees, and cucurbits are important sources of pollen and nectar during their blooming periods.

Other plants such as wild buckwheat, star thistle, sage, and fireweed in restricted localities may yield commercial quantities of honey in favorable years and may rank high in the estimation of the beekeeper because of their value to the bees as a source of food for building up the colony early in the spring or to carry it over the winter period.

The region varies in rainfall from 1 to 2 inches in the desert areas to more than 60 inches in the rain-forest area, in elevation from below sea level to snow-capped mountains, and in temperatures from dry and hot to humid and extremely cold.

Migratory beekeeping is practiced by most of the commercial beekeepers, and four or more moves per year are not uncommon. In California, most of the bees are held in almond areas during the winter. The almond orchards are distributed from Chico in Butt County in the north to Kern County in the south. The pollination season begins with almond blooms in early February. As the almonds finish blooming, the plums and prunes begin to bloom and the beekeeper may move to these. Cherries bloom near mid-March through mid-April. After this period of fruit bloom, there is a dearth of pollen and nectar in cultivated areas. To maintain and build up colonies for summer pollination service, the beekeeper moves his bees to the mountains where bees are held in manzanita and sage at elevations around 2,000 to 6,000 feet.

Native plants supply pollen and nectar in the Sierra Nevada range, the coast ranges, and coastal areas between the Pacific Ocean and the coast ranges.

Commercial pollination service begins again in June-July with melon pollination, ladino clover, and alfalfa seed production. In the fall, after these sources have been harvested, the beekeeper moves his bees into native flora along the east side of the coast range. This is a major source of nectar and pollen for winter stores from August to frost.

California beekeepers south of the Tehachapi Mountains begin to build up their colonies on native plants in January. Until citrus bloom in April, this is the main bee pasture. Some southern California beekeepers move into the southern and central almond areas in February and March and into alfalfa and cotton during the summer and early fall.

It is evident, therefore, that the beekeeper must move his bees to take advantage of pastures offered by native and cultivated plants during the period. The placement of 2,000 colonies from several beekeepers in a solid square mile of alfalfa grown for seed is not unusual. The use of bees for pollination is extensive. An estimated one-half or more of all colonies are used sometime during the year for pollination hire.

In the last few years, many beekeepers have had to replace almost 100 percent of their colonies due to pesticide losses. These losses are increasing each year. The major dollar loss to beekeeping in California is caused by (1) pesticides, (2) wax moth, and (3) foulbrood diseases.

Beekeepers operate an average of 2,000 colonies. In such operations, the apiary rather th