The Other Side of Beekeeping - September 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Family Liliaceae--The Lily Family

   The story of the origin of hybrid onions is in my opinion quite interesting since nearly all the onions we see on the grocery shelf today rely on one onion found back in 1925.

Depending on the text consulted, the Liliaceae consists of about 240 genera and between 3000 and 4000 species. One of the major characteristics of the Liliaceae is that they are all monocots. The true flowering plants (angiosperms) were once divided into two subgroups, which most of us learned as the monocots and the dicots. This has now changed into the monocots and the eudicots (the true dicots) and a small group that doesn’t fit well into the eudicot definition. This last group doesn’t appear to be of much importance to the beekeeper, and for the purposes of this discussion, we will deal with only the monocots and eudicots. The monocots have only one seed leaf (cotyledon) compared to two in the eudicots. This is the first green leaf to emerge from the seed and contains sugars and other nutrients that are used by the plant during its initial stages of growth. In addition to the single cotyledon, the monocots have their vascular transporting system (xylem and phloem) located throughout the stem rather than in the periphery of the stem as in the eudicots. The first root to emerge during the germination of a monocot seed generally dies, and there is, therefore, no tap root system as is common in the eudicots. Instead, the “roots” (called adventitious roots) are derived from the underground part of the plant, which is mainly stem tissue, not true root tissue. The pollen grains of monocots have only one pore or furrow while those of the eudicots have three. The flowering parts are usually in threes (three petals, three sepals etc) compared to the usual fours and fives in the eudicots. The leaves of monocots are generally elongated and have parallel veins. Be aware that there are some exceptions to these generalizations. While the monocots make up only about 25% of the angiosperms, some of the monocots are important to beekeepers and include the agaves, skunk cabbages, palms, lilies and the grasses and their kin (example corn)[24 & 31].
The Liliaceae is generally distributed over the earth, but is most abundant in the temperate and subtropical regions. Members of the family are mostly perennial herbs, which after the seedling stage, form food storage and reproductive organs: bulbs, corms, rhizomes and occasionally tubers1.

The leaves, which only rarely persist throughout the year, emanate either from the underground storage structure or from the stem, in which case they are usually placed alternately or whorled around the stem and are only rarely placed oppositely. The leaves are typically elongate and narrow and parallel-veined, but occasionally are broad and net veined as in the trilliums.

The flowers are usually bisexual, radially symmetrical, often showy, and arranged in various types of inflorescence. The tepals2 are of similar size, shape and orientation and usually occur in groups of three, less often in groups of two or four, and there are generally as many stamens as tepals. There may also be a corona3.

The ovary placement ranges anywhere from superior to inferior4 and generally has three cavities (locules) in which the several to many ovules later become seeds. Each ovule is attached to the central axis of the ovary (axial placentation). Nectaries are often located on the outer tepals. The fruit is a berry or a capsule5 with few to many seeds.

The Other Side of Beekeeping - August 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

More Members of the Ericaceae

Scientific name: Gaultheria shallon

Origin: Gaultheria shallon is native to the Pacific Coastal Area of North America from California to Alaska.

Plant description: Salal is a low-spreading shrub often with erect stems that can grow to heights of about 6 ft. The leaves are 2 to 4 inches long. They are quite variable in shape and range from being kidney-shaped and strongly indented at their base (reniform) to being nearly squared off at the base, then often being 3 to 4 times longer than wide with more or less parallel sides (oblong), to being widest in the attached half (ovate), in which case they can end in a sharp point, or be more rounded (orbicular)1. The leaf edges are frequently finely toothed (serrate). The individual flowers are borne on glandular hairy racemes2. The urn-shaped flowers range in color from white to pinkish and are about 0.375 inches long and are also frequently also glandular-hairy. The more or less spherical fruit generally has a diameter of 0.28 to 0.3 inches and is dark purple, later becoming black.[5, 13, 16, 25]

Distribution: In California the species generally inhabits moist forest margins under about 2600 ft and prefers acidic soils[25] (pH 5.5-7.0[24]). Goltz[8] states “Salal is the most abundant shrub in open timbered areas and thickly covers the forest floor in western Oregon. It yields some honey on the west side of the Cascade Mountains in Oregon where its growth is less rank.”

Blooming period: According to Hortus Third[10], salal blooms spring to early summer. In California it blooms during April to July[16]. In Oregon it blooms from May into July[5]. Ramsay[20] provides a blooming date range of June to mid July for Canada. D. M. McCutcheon in his response to the Ayers and Harman questionnaires[3] indicates that it blooms during June and July in British Columbia. Sheppard et al.[22] indicate that it blooms during May and June in British Columbia.

Importance as a honey plant: Ramsay[20] provides the information that salal is “regarded as one of the best native nectar plants in western Canada, especially coastal British Columbia.” McCutcheon in his reply to the Ayers and Harman questionnaire[3] considered the species to be a major source of nectar in the mountainous area of British Columbia. Sheppard et al.[23] lists the species as being among the “Principal nectar-bearing native flora of British Columbia’.

The Other Side of Beekeeping - July 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

Cranberry

Cranberry
Scientific name: Vaccinium macrocarpon
Synonyms: Oxycoccus macrocarpus
Origin: Northeastern North America

Plant description: Cranberry is an evergreen or semievergreen creeping, mat-forming, freely rooting trailing vine, that spreads to about 3 ft across and sends up numerous vertical branches known in the trade as uprights that reach heights of 6 to 18 inches. Both the trailing vines and the uprights have leaves, but only the uprights produce fruit[11]. The oblong, elliptic leaves range in length from about 0.33 to 0.75 inches and are somewhat whitish beneath. The usual 5-6 flowers per upright range between 0.25 and 0.33 inches in length and are arranged in small lateral clusters. At first the blossoms are white, but if not pollinated remain on the upright and become a rosy pink. The stamens fit tightly together and form a tube around the pistil. At first the stigma is retained within the staminal tube, but as the style lengthens, it is pushed through the opening at the free end of the tube. The pollen from each stamen is released through a small opening called a pore that is located at the stamen's terminal end. The nectaries are found surrounding the style just inside the ring stamens. The flowers hang with the tips of the stamens pointed downward and the petals curved upward. Some think the flower in silhouette resembles the neck and head of a crane. The species was originally called craneberry and subsequently shortened to cranberry. The roundish red fruits range in diameter from about 0.25 to 0.75 inches[16 &21].

Distribution: In the wild, cranberry is found in acid bogs and swamps with a pH generally between about 3.2 and 4.5[16]. In the US, the states in order of descending production are WI, MA, NJ, OR, WA[33]. In Canada, the major provincial production occurs in British Columbia. Other areas of Canadian production occur in Quebec, New Brunswick, Nova Scotia and Prince Edward Island[34].

Blooming period: In Massachusetts the plant blooms in late June and early July and the fruit ripens in September and October[11]. In Oregon, Wisconsin, and British Columbia the species generally blooms during June and July[35].

Importance as a honey plant: While honey is sometimes obtained from cranberry, it is not a major honey producer, but see below under ‘Honey'. Ayers and Harman[1] found the species to be listed as a source of commercial pollination in WA, OR, WI, NJ, and MA and in the Canadian provinces of BC and NB.
 
Honey potential: Shaw et al[24]found that the nectar sugar concentration of cranberry varied between 38 and 62 %. The concentration tended to increase as the daily temperature rose. They also found concentration differences between the three cultivars with which they worked. The variety MacFarlin produced a lower average nectar sugar concentration (45.7%) than either Howes (50.2%) or Early Black (54.7%). Singh[27] in India, found the sugar concentration of cranberry nectar ranged between 16 and 51%.

Honey: Irving Sibert[26] describes cranberry honey obtained by Justin Caswell (an early cranberry pollinator) as, "The honey is reddish in hue and has a spicy flavor. I think it is as good as any I've ever tasted and it has a flavor all its own." He quotes Caswell as, "Folks pretty near stand at the extractor waiting to buy it." Caswell himself states, "Honey from cranberry is a light to medium red, of mild flavor and not as sweet to the taste as some honeys[3]." Gates[13] (1918) stated that cranberry produces a superior honey. McGregor[21] also reports that cranberry growers sometimes produce a reddish honey they associate with cranberry. White[31] analyzed two quite similar cranberry honeys that were submitted by different beekeepers located in Eastern Massachusetts (See table 1).

Pollen: Shimanuki et al,[25] describe separating cranberry pollen from other pollens in pollen traps that came from hives near a cranberry bog based on its "yellowish brown color".

Additional Information:
Early history of the American cranberry industry
The American cranberry industry initially started along the Cape Cod and New Jersey coasts in about the 1830s to provide sailors sailing out of Boston, New York and Philadelphia with a long-lasting food supplement that prevented the disease scurvy that caused devastating health problems to sailors of the day[28]. Today we know that the active agent was, of course, vitamin C.

How the flower and bees function
Roberts and Struckmeyer[22], working in 1942 in Wisconsin, stated that the bee did not touch the stigma when visiting cranberry and, therefore, pollination resulted from a combination of the bee jarring the flower and dispersing the pollen into the air, which the wind then distributed to mature stigmas. They presented a small amount of data that suggested brushing the uprights with a "stick or paddle" increased berry set (their table 2). As a result of this assertion there were cranberry growers in Wisconsin who actually dragged heavy ropes over their plants to simulate these actions. These ideas have largely been discarded today, and there is now considerable evidence that insects, especially bees of various types, are largely responsible for pollination[18]. When the flower opens, the style is slightly shorter than the ring of stamens in which it is encased. About a day before the style lengthens and the stigma is pushed out of the staminal tube, the anthers release their pollen through the openings (pores) at the end of each stamen. When the pollen is shed, the stigma is dry. The stigma remains unreceptive for about 24 to 36 hrs after the anthers initiate their pollen release. By this time the style has lengthened to place its now moist, sticky and receptive stigma about 1/16 inch below the openings of the now empty anthers[21]. It is now conceded that the pollen is not windblown and is not likely to come in contact with its own receptive stigma. The pollen grain is made up of four pieces (a tetrad), each capable of germinating into a pollen tube and fertilizing an ovule and, therefore, relatively small amounts of pollen are needed to fully pollinate the flower[22].
When a bee probes an early stage flower (before the stigma is receptive), the pollen falls out through the anther pores onto the bee's body[12]. When the bee then moves to a more advanced flower, it transfers that pollen to the now moist, sticky, and receptive stigma, causing cross pollination. Cranberry thus appears to be designed to facilitate (perhaps force is a more appropriate word) cross-pollination. After a flower is pollinated, it sheds its petals and initiates fruit development. If the flower is not pollinated, it persists for some time on the plant and the petals turn a rosy color. A high incidence of rosy colored flowers is one of the keys to identifying inadequate pollination[21], but see the discussion below about blasting.
In addition to facilitating fruit-set, bees also help increase the size and uniformity of the berry. Each cranberry consists of four basic female units (carpels) which contain several ovules, each capable of developing into a seed if fertilized. Filmer et al. [9], using four different cranberry varieties, demonstrated quite convincingly that berry size is positively correlated with the number of seeds/berry. In addition to being small, berries with low seed numbers are also often misshapen.

Which bees are the best pollinators?
Originally, as the cranberry industry developed along the North American East Coast, native pollinator populations were apparently adequate for the pollination of cranberry bogs. In a 1914 extensive report on cranberry research[10], under the subheading ‘Blossom Pollination', while there is a clear indication that the author was aware of "bees" being important in setting cranberry fruit, the words "honey bee" aren't mentioned. In 1940, a time when Massachusetts was claimed to produce more than half of the world's cranberries, another extensive report entitled ‘Cranberry Growing in Massachusetts' doesn't seem to mention pollination at all, much less honey bees. It is as though pollination just magically occurred, and under the environmental conditions of the day that provided adequate native bee populations for pollination, that's probably about the way it appeared.
Today native bees frequently aren't available in sufficient populations to effect good cranberry pollination. As an example, Winston and Graf[32] found the native bee populations in The Fraser Valley of British Columbia to be very low and insufficient for pollination of several types of berries, including cranberry. The reasons appeared to include: pesticide impact, habitat destruction, competition with managed honey bees and an extended rainy period during the spring in which the study was done, which may have washed out nesting sites1. In addition to these reasons, there are numerous suggestions in the literature that as civilization encroaches on cranberry bogs, there are other reasons that the soil-nesting habitat of bumble bees is deteriorated (soil compaction, pavement, shifts in vegetation type, etc.). As it became clear that native pollinator populations were diminishing to the point that it was adversely affecting cranberry pollination, the interest in honey bees for cranberry pollination grew. I find ironic, that as early as 1925 Ray Hutson, one of the early proponents of honey bee pollination of cranberry, pointed out that there were many native pollinators available in cranberry bogs, which he attributed to the undisturbed space surrounding the bogs that he called "waste land"[14].
In reality, honey bees are not very fond of cranberry, and on a bee per bee basis, honey bees are not nearly as effective as bumblebees for cranberry pollination (see ‘pollination recommendations' below). Cranberry produces only small amounts of nectar, sometimes almost none, and is also frequently considered to be at best a marginal pollen producer2[4 & 17]. When compared to bumble bees, the main advantage of honey bees is that they are relatively easy to manage. Overnight the cranberry grower can have multiple hives of honey bees delivered, each of which may have a larger population than the total population of local native pollinators. While honey bees are not the best cranberry pollinators, there is considerable evidence that they can, under the right circumstances, effect good cranberry pollination. As early as 1925, Ray Hutson[14] , working in New Jersey, caged equal areas of a cranberry bog with and without bees. In the cage with bees there were 2385 flowers that produced 1335 berries (a 56% set), while in the cage without bees, 2184 flowers produced 185 berries (an 8.5% set). In 1947 Farrar and Bain[5 & 6] in Wisconsin found only 10 berries/ft2 from caged plants without bees, 124 berries/ft2 from open pollinated plants (no cages) 3 and 171 berries/ft2 from plants caged with bees. Filmer and Doehlert[7 & 8] reported unreferenced New Jersey cranberry research, which was apparently designed to refute the Roberts and Struckmeyer work described above. This New Jersey research produced only 15 cranberries/ ft2 in cages where bees were excluded, even though the vines were agitated daily by various means to dislodge pollen. In comparison, 90 to 152 berries/ ft2 were produced in adjacent uncaged plots where apparently pollinators were plentiful.
Even when honey bees are plentiful, they don't always do a good job of cranberry pollination. Kevan et al.[15], for example, in 1983, working in Ontario, studied the effect of honey bees on pollination using the relationship between distance from the hive and fruit set4. These researchers found no significant differences between distance from the hives in either fruits/flower, and seeds/flower even though honey bee populations did decrease with distance from the hives. Even near the hives, however, the population of foraging honey bees was not high. There were, however, apparently relatively large populations of bumble bees in the area. It is clear that cranberries are not very attractive to honey bees, and if they have another and better foraging choice, they will take advantage of it. In this study it appeared that the honey bees were working more attractive plants in the area, and when it came to cranberry pollination, they were leaving the heavy lifting to the bumble bees. Ironically, if for some reason that particular year, the area surrounding the bog hadn't supported bumble bee populations, that space, so important to maintaining bumblebee populations, might well have turned around and "bitten" the cranberry grower.
Marucci (1967)[17], reviewing some of the New Jersey research, reported even strong honey bee colonies did not do much pollinating until about the 7th to 10th day after cranberry had started blooming. In part this was again because there were numerous competing higher quality forages in the area that were terminating bloom just as cranberry began to bloom. In this review, the author describes some of his unpublished data where he had manipulated small cages in a cranberry bog to study the effect of various pollinator exclusion periods on berry production. He found that the percentage fruit formation was not reduced by one or two or even sometimes three weeks of pollinator exclusion, if one week of unhampered foraging had been allowed during peak bloom. If, however, the one week uncaged period occurred at the beginning of bloom, fruit set was greatly reduced[17].
The work of Shimanuki et al.[25] suggests that bees should be moved into the cranberry bog before the peak of bloom. In this work, three hives were moved into cranberry approximately one week before peak bloom, and another three hives were placed there at peak bloom. The hives placed there before peak bloom produced 87.9 grams of cranberry pollen (74.25% of pollen collected) versus 15.77 grams of cranberry pollen (20.94% of pollen collected) by those moved into the bog at peak bloom. Notice that this may seem a little different than the pollination advice that is sometimes given, i.e., that bees should not be moved to a pollination site too early or they may become "addicted" to the surrounding bee forage and continue to work it instead of the crop for which they were intended. While this may seem different, it may not be, given the relatively long blooming period of cranberry, which is about 4 weeks[17].

Blasts
As described above, unlike other deciduous fruits, unpollinated cranberry flowers remain on the plant and turn a rosy red. In this condition they are called "blasts". Generally many blasts can be seen in a cranberry bog, and are frequently of concern to the cranberry grower. Marucci and Filmer[20] compared fruit set on cranberries caged with a hive of bees, which they considered represented an overabundance of bees, versus uncaged plants exposed to a honeybee population of one hive/acre. The caged plants with bees did not set a higher percentage of fruit than the uncaged plants. In their experiments they also pruned flowers from uprights and counted the subsequent number of blasts. Pruning reduced the number of blasts and gave a higher percentage of flowers that produced fruit. They also noted that "pruning of florets" by frosts gave the same result. Their final conclusion was that an insufficient number of bees would increase blasting, but even with an overabundance of bees, relatively high rates of blasting would still occur. In this view, honey bees can minimize the number of blasts by providing the pollination necessary to produce the maximum number of fruits that the plant can sustain, but blasting is rarely reduced to much below 50% in New Jersey where their study was done[17].. This work also demonstrated pretty conclusively, up to a point, that cranberry production per acre was directly related to the number of uprights. This relationship, however, can't extend to an infinite number of uprights. Whereas Marucci and Filmer's[20] data ends at about 200 uprights/ft2, the data of Roberts and Struckmeyer[22] extends to over 500 uprights/ft2 and in their study the production dropped off fairly precipitously at about 250-260 uprights/ft2.

Pollination recommendations
Honey bee recommendations
In his review of the pollination recommendations given for cranberry, McGregor[21] stated, "The pollination recommendations for cranberries lean consistently toward the use of more colonies of honey bees per acre". Perhaps this is partly because of the deterioration of native pollinator populations over time. He provides the data found in Table 2. Scott-Dupree (1995) recommended 1 colony per acre for Canadian cranberry producers[23]. Delaplane and Mayer[4] (2000), after reviewing the literature, state that the average literature recommendation rate is 3 colonies/acre.

Recommendations for Other pollinators
Hutson[14] recommended 448 bumble bees per acre. This was based on the fact that there were that many bumblebees in a cranberry bog that was relatively devoid of other insects, but set a good commercial crop.
Cane et al.[2l] estimated the number of leaf-cutting bees (Megachile addenda) needed to produce a commercial crop of cranberries by two methods: (1) by counting the number of pollen grains removed per flower and comparing that figure to the number of pollen grains in completed nest cells5 to estimate the number of flowers visited and (2) by using the floral visitation rate, foraging trip duration and the number of trips needed to complete a nest cell. The two estimates of the number of flowers visited per nest cell were exceptionally close (1076 and 1207, respectively). In good foraging weather they estimated that 451 nesting M. addenda females per acre would be sufficient to provide a commercial harvest. The authors point out, however, that they encountered high rates of nest parasitism from the cleptoparasite6, Coelioxys immaculata, also in the same family (Megachilidae) as its host, M. addenda. This parasitic species would have to be controlled if M. addenda were to be relied upon for pollination.
Delaplane and Mayer[4] point out that given the potential of bumble bees for pollination, it would seem advantageous for cranberry growers to manage the area around their cranberry bogs to encourage higher populations of these organisms. They point out that conceivably this could be done by (1) leaving the land around the bogs undisturbed to encourage nesting sites; (2) providing artificial nest boxes along the edges of the bog; (3) culturing supplemental bee pasturage in the areas around the cranberry bogs to promote colony health after the cranberry flowering season. In general, however, such bumble bee conservation measures in cranberry have apparently not been measurably successful.

Potential for developing a cranberry honey bee.
Shimanuki et al.[25] noticed one of the hives used in their pollination timing study described above consistently produced significantly more cranberry pollen than the other hives (see table 3). Apparently, queen selections from this hive were made that produced colonies that were better than average cranberry pollen collectors, but they were so vicious that they were destroyed[28]. This finding does, however, suggest that it would be possible to breed bees better adapted to cranberry pollination.

Potentials for cross pollination
Marucci and Filmer [19] reported that it had been observed that sometimes cranberry bogs with mixed varieties had greater cranberry productions than nearby bogs with only one variety. These greater productions were sometimes as much as five times the state average. An experiment was set up where mixed varieties were caged with bees in such a way that the bees could work both inside and outside the cages. This caging arrangement was intended to supply approximately equal bee populations inside and outside the cages. There were also two bogs sampled that contained intermingled plants of different varieties that were also sampled. The results were promising, but Free[12] points out that it is not clear that these outcomes did not result from larger bee populations in the areas with mixed varieties than in the areas with a single variety, and that more research should be done.

Colony deterioration in cranberry bogs
Marucci[17] reported that there is an unusually high incidence of European foulbrood, as well as both morale and colony size deterioration, when honey bees are set out in New Jersey cranberry areas. Interestingly, because cranberry and blueberry belong to the same plant family (Ericaceae), a similar situation seems to occur in Michigan when bees are used for blueberry pollination[30]. Perhaps this is because the environments are to some extent similar (low lying areas, with acidic, high organic soils).

References  
1. Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In The Hive and the Honey Bee (J. M. Graham, Ed.),  Dadant and Sons. Hamilton, IL.
2. Cane, J. H., D. Schiffhauer and L. J. Kervin 1996. Pollination, foraging, and nesting ecology of the leaf-cutting bee Megachile (Delomegachile) addendo (Hymenoptera: Megachilidae) on cranberry beds. Annals of the Entomological Society of America 89:361-367.
3. Caswell. J. H. 1962. Caswell Bee Company-Cape Cod cranberry pollinators. American Bee Journal 102:222-223.
4. Delaplane, K. S. and D. E. Mayer 2000. Crop Pollination by Bees. CABI Publishing. New York.
5. Farrar, C. L. and H. F. Bain. 1946. Honey bees as pollinators of the cranberry. American Bee Journal86: 503-504.
6. Farrar, C.L. and H. F. Bain. 1947. Honeybees as pollinators of cranberries. Cranberries 11(9):6-7, 22-23
7. Filmer, R. S. and C. A. Doehlert. 1952. Use of honeybees in cranberry bogs. New Jersey Agricultural Experiment Station Bulletin 764. 4 pages
8. Filmer, R. S. and C. A. Doehlert. 1959. Use of honeybees in cranberry bogs. New Jersey Agricultural Experiment Station Circular 588.
9. Filmer, R. S., P. E. Marucci and H. Moulter. 1958. Seed counts and size of cranberries. American Cranberry Growers' Association Proceedings 88:22, 26-30.
10. Franklin, H. J. 1914. Reports on experimental work in Connection with Cranberries. Massachusetts Agricultural Experiment Station Bulletin 150:37-62.
11. Franklin, H. J. 1940. Cranberry growing in Massachusetts. Massachusetts Agricultural Experiment Station Bulletin 371.
12. Free, J. B. 1993. Insect Pollination of Crops. Academic Press. London.
13. Gates, B. N. 1911. The honey bee and cranberry growing. Cape Cod Cranberry Growers' Association. Annual Report 24:28-29.
14. Hutson, R. 1925. The honey bee as an agent in the pollination of pears, apples and cranberries. Journal of  Economic Entomology 18: 387-391.
15. Kevan, P. G., R. M. Gadawski, S. D. Devan and S. E. Gadawski. 1983. Pollination of cranberries, Vaccinium  macrocarpon, on Cultivated marshes in Ontario. Proceedings of the Entomological Society of Ontario 114:45-53.
16. Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dectionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co., Inc. New York.
17. Marucci, P. E. 1967. Cranberry Pollination. American Bee Journal 107: 212-213.
18. Marucci, P. E. and H. J. Moulter. 1977. Cranberry pollination in New Jersey. Acta Horticulturae 61:217-222.
19.Marucci, P. E. and R. S. Filmer. 1964. Preliminary cross pollination tests on cranberries. American Cranberry Growers' Association Proceedings 91st-94th Annual Meeting. 1961-64:48-51.
20. Marucci, P. I. and R. S. Filner. 1957. Cranberry blossom blast is not caused by disease New Jersey Agriculture 39: 8-9.
21. McGregor. S. E. 1976. Insect Pollination of Cultivated Crop Plants. Agriculture Handbook No. 496, Agricultural Research Service. United States Department of Agriculture. Washington D.C. This publication is being updated and is available on the web at gears.tucson.ars.ag.gov/book
22. Roberts, R. H. and B. E. Struckmeyer. 1942. Growth and fruiting of the cranberry. Proceedings American Society for Horticultural Science 40: 373-379.
23. Scott-Dupree, C. M. Winston, G. Hergert S. C. Jay, D. Nelson J. Gates, B. Termeer and G. Otis (Eds.) 1995. A Guide to Managing Bees for Crop Pollination. Canadian Association of Profesional Apiculturists.
24. Shaw, F. R. , W. M Shaw and J. Weidhass. 1958. Observations on sugar concentrations of cranberry nectar. Gleanings in Bee Culture 84:150-151
25. Shimanuki, H., T. Lehnert, and M. Stricker. 1967. Differential collection of cranberry pollen by honey bees. Journal of Economic Entomology 60: 1031-1033.
26. Sibert, I. 1967. The cranberry pollinator. Gleanings in Bee Culture 95:281-282.
27. Singh, S. 1954. Insect pollinators and the breeding of fruit varieties. Indian Journal of Horticulture11:6-9.
28. Stewart, J. D. 1970. Cranberry Pollination in New Jersey. In: The Indispensable Pollinators. Arkansas Agricultural Extension Service Miscellaneous Publication 127:181-184.
29. USDA, NRCS. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA
30. Wardell, G. I. 1982. European Foulbrood: Association with Michigan blueberry pollination, and control Ph.D Thesis. Michigan State University, Department of Entomology.
31. White, J. W., M. L. Riethof, M. H. Subers and L. Kushuir.  1962. Composition of American Honeys. U.S.D.A. Technical Bulletin No 1261. U. S. Government Printing Office. Washington, D.C.
32. Winston, M. L. and L. H. Graf 1982. Native bee pollinatiors of berry crops in the Fraser Valley of British Columbia. Journal of the Entomological Society British Columbia 79:14-20.
33. http://usda.mannlib.cornell.edu/usda/current/Cran/Cran-08-18-2009.pdf
34. http://www4.agr.gc.ca/AAFC-AAC/display-afficher.do?
id=1241547089433&lang=eng#s3
35. From the the original questionnaires used in reference 1

The Other Side of Beekeeping - June 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

The Balsaminaceae
The Touch-me-not Family

Depending on the reference consulted, the Balsaminaceae is  represented by 2-4 genera and between 450-850 species of  mainly herbs and subshrubs primarily from Eurasia, North America and Africa. Part of the family's largest genus, Impatiens, is native to North America. The USDA website[20] lists 11 species of impatiens growing in North America, 5 of which are considered to be native.

The Family is characterized by crisp, translucent, watery stems that have a glassy appearance. Frequently the stems are tinged with red and purple and have swollen nodes. The leaves are usually arranged on their stems alternately, but may have opposite or whorled arrangements as well. Usually there are no stipules1.

The flowers are complete, bilaterally symmetrical, sometimes referred to as highly irregular2 and are either arranged singly or with several flowers coming from a common floral stem.
The calyx has 3, rarely 5, often colored sepals, the lower one frequently extended posteriorly into a funnel-like nectariferous spur3. The corolla usually consists of 5 petals, but sometimes 4 or 2. The upper petal is generally free (not joined to other petals), flat or helmet shaped, and the four lower and side petals are usually joined in pairs, one pair on each side of the flower.

The five stamens have short filaments and the anthers are joined to form a cap over the pistil. The style of the pistil is very short. As the ovary enlarges, the pistil breaks through the cap. The stamens, petals and sepals are attached to the floral stem below the ovary (ovary in superior position). The pistil is compound with five carpels4, and there may be one or five stigmas.

The generally elongate fruits have five sides (valves), which, when ripe, split apart and explosively scatter their seeds when touched, therein the origin of the common name, "touch-me-not. Rarely the fruit is a berry5.

About 20 species of the genus Impatiens are cultivated. [2, 4, 5, 7, 11 & 19].

The Other Side of Beekeeping - May 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

Soybeans Continued

In the April column I discussed the slow historical acceptance that soybeans can sometimes provide a honey crop, sometimes even a good honey crop. It is clear from the data that was provided that there is a wide range of innate honey potentials within different soybean lines. That's not the end of the story, however. It may not even be the most important part of the story. The local environment is also very important in determining the honey potentials of the soybeans grown in a particular area.

Environmental effects on soybean honey production
 In 1979 Erickson and Robins[10] undertook what I consider one of the first steps in a string of important studies to definitively identify the environmental factors that lead to high soybean honey production. In this early study they set out to test the hypothesis that historic soybean production records, as well as soil type based on existing soil maps would be correlated with honey production. If that were true, soybean production records and/or soil maps could be used to locate apiaries that were likely to provide good honey production. The study was done in the Mississippi delta areas of Missouri and Arkansas where the Ohio and Mississippi Rivers, as well as several smaller tributaries, have changed courses several times in recent geological history, leaving a variety of soil types that made it possible to test this hypothesis. In general, the study confirmed the original hypotheses, but honey production was more highly correlated with soybean production than with soil type. The soil characteristics that correlated with good honey production were deep, heavier, highly fertile soils (including potassium), with a pH between 6.0 and 6.41. As final proof, in the authors' words, "Relocation of apiaries short distances (up to 10 miles) to heavier soil and class sites invariably improved the productivity of the colonies involved." Their conclusion was, "Whether or not the nectar source is soybeans, beekeepers seeking productive apiary sites should, in lieu of more definitive information, do best by locating on those farm lands with the best records for crop productivity (yield/acre)."
 The Erickson and Robins study confirmed the more casual observations of earlier authors. Pellett[20] reports from a 1922 letter sent to him by a Mr. J. R. Pinkham of Washington NC (Coastal area) that soybeans do not seem to yield as heavily on uplands as on the black swamp or Pocosin silt. Davis[3] stated, "The best soybean nectar-producing areas of Arkansas are the river bottoms where the soil is deep and fertile." Harvey Lovell[17] reported that a beekeeper near Geneva, NY found that bees worked soybeans heavily and made some honey from soybeans growing on light gravelly soils, but made no honey when grown on clay.
 With the above study as a background, the research group (Robacker et al.[22]) went on to study the effects of other environmental factors on soybean honey production in a more controlled way in the University of Wisconsin's Biotron where factors such as day and night air and soil temperatures, nitrogen, phosphorous, potassium and other soil nutrients, light intensity, soil moisture, etc. could be managed alone or in various combinations [22]. Over 50 combinations were used and their effects on the soybean recorded. They looked at plant size, flowering date, number of flowers produced, flower color intensity, degree of flower openness, nectary development, nectar secretion, plant aroma, and attractiveness to bees. It was easy to optimize one parameter at a time, daytime temperatures, for example, where all the other variables were held constant. It soon became apparent, however, that this daytime optimum temperature held only for that particular set of other conditions. For example, using primary plant characteristic such as growth, relatively warm nighttime temperatures could offset the effect of what had originally appeared to be nonoptimum daytime temperatures. As another example, soil nitrogen levels affected optimum day/nighttime temperature regimes. They, in fact, found that soil nitrogen and phosphorous levels affected almost everything else including themselves. These interrelationships are called interactions. They sound messy, and they are, but this is the way the world works. One of the prime axioms of System Science is that you can't optimize all the variables of a complex system at the same time. As messy as I just made it sound, a lot of good information and some very interesting questions and theories grew out of this study. First it began to make clear why the observations made by beekeepers often differed. It also began to clarify which plant characteristics were important in attracting bees and which environmental elements affected these characteristics. Fig. 1 represents the authors' attempt at identifying and assigning relative values to the soybean characteristics that affect the attractiveness of soybeans to honey bees. Some of these are quite obvious, for example flower openness-if a flower doesn't open, it probably won't be attractive to bees. Notice that this list is also not without its obvious interactions. If, for example, flowers don't open, the number of flowers probably isn't going to greatly affect the attractiveness to bees.2

Effect of temperature
 The researchers found that warm day temperatures (about 83˚F) provided the best results. Warm nighttime temperatures also seemed to contribute to attractiveness. This study also confirmed the earlier more casual observations of others. The heavy soybean flows of both Johnson[14] and Milum[19] (see ‘Honey Potential, April Column) occurred after rains that were followed by warm temperatures that reached 100oF in the Johnson situation, and temperatures, while warm in the Milum situation, were a little cooler than those experienced by Johnson3. Milum also tells of communication with a Mr. Kirk from Farmersville, IL (ca. 90 miles southwest of Urbana) who had also experienced a soybean honey flow during the same hot, dry weather system. Milum concluded that adequate moisture coupled with hot, dry weather may be one of the factors controlling soybean honey production. Jaycox[13] states, "I have found that our scale colonies usually gain more weight during soybean bloom if the temperatures are consistently in the 80's or above." Erickson had also previously reported adverse effects of cold weather on soybean honey production. In a three-year study [6 & 7]4 using many varieties, he noted great variation in the effect of temperature on flower openness and nectar secretion between those varieties. Many of the varieties didn't secrete any nectar. He found that one variety, ‘Hark', which was less adversely affected by cool temperature than most varieties, ceased producing nectar and the flowers remained closed (cleistogamy) at mean daily temperatures below 70oF (21oC), and one to four days of warmer temperatures were necessary to again stimulate nectar secretion. Interestingly the nectar sugar concentration of those flowers that produced nectar seemed to remain relatively constant, between 33 and 36%5 during these up and down nectar production periods.

Soil fertility
 In the Robacker et al. Biotron study[22], high nitrogen and low phosphorous situations also led to high levels of attractiveness to bees. Soil temperatures, and surprisingly, the two factors, potassium, and soil moisture had little effect on attractiveness. The authors point out, however, that soil moisture levels were the hardest condition to maintain in the Biotron.

The Aromas
 In a 1982 Robacker et al. article[ 22] the authors describe two aromas. This was expanded to three in another article by Roebacker et al.[23]. Here I use the three-aroma version, and because there are several things going on at once, the reader is encouraged to follow the associated graphic as they read what is provided in the text. A soybean flower is open for only a day. Early in that day, at a time when the flowers were not yet open, they released a mixture of aromas where component (1) predominated. As the flowers opened, nectar secretion started and reached a maximum at about 3.5 hours into the light cycle, the same time aroma (3) reached its maximum, and aroma (1) and (2) reached their low points. A little later as the nectar secretion diminished, the concentration of component 3 dropped, and by the time nectar secretion ceased at 8 hrs, aroma (1) had again become the major component. Thereafter, as the flower closed, the concentration of all aroma components diminished, with component (1) still predominating. Notice that the bee foraging population reached its peak as component (3) reached its peak. Through hours 5.0 and 6.5 the foraging population remained high because there was still some nectar and probably also pollen that could be collected. Then, as nectar secretion continued to diminish, component (1) regained its original prominence and the foraging populations declined. Recalling that bees have a memory and can associate "cues" with recurring events, in this case, associating aromas with nectar production, it is almost as though the plant were telling the bees "not yet" then a little later "we're ready for you now" and still later "it's too late now, look for another flower". While that's interesting, plants aren't altruistic; they wouldn't be trying to help the bees for the bees' sake. The researchers speculated that this seemingly altruistic behavior ensures that both self and cross-pollination will occur. Using what seems like a reasonable train of logic, they estimated that it would take about 20 pollen grains to fully fertilize the average soybean flower. The top graph of the associated figure represents, over time, the percentage of stigmas that would have 20 or more pollen grains (full self-pollination) in a bee-free environment. Up until about the time that the bees are receiving some nectar and their foraging populations begin to increase, mainly self-fertilization would occur. As the foraging bee population increases, the process of cross-pollination would be initiated and increase as the foraging population increases. Notice that the foraging population reaches its maximum at a time when there would be, without bees, only about 33% of the stigmas fully self-pollinated (have at least 20 pollen grains). Some self-pollination would also be occurring at this time as a result of the bees cavorting around on the flower. The strategy seems designed to enforce a balance between self and cross-pollination.

Interesting questions
 The Robacker et al. article[22] described above addresses some very interesting theoretical questions. Why would a plant species known to frequently self-pollinate before it opens "be interested" in attracting bees with contrasting coloration, nectar guides, and nectar production etc. (see April column). The researchers provide an interesting answer. First, they explain that being either totally cross-pollinated or totally self-pollinated probably isn't good, and plants evolve striking a happy medium between the two strategies that fits the environment in which they find themselves. Then, they propose the hypothesis that the progenitor of the current soybean was originally a largely cross-pollinating species, but when it was moved to many parts of the world as an agricultural crop, it experienced environments devoid of its original pollinators and the "tug of war" between self-pollination and cross-pollination slid the soybean toward self-pollination. In this view, many of the attributes of a cross-pollinating plant have not yet been lost (the showy floral display, nectar, nectar guides, the opening to the tongue channel and the associated tongue guides, attractive aromas, etc.6). Whatever the explanation, it will be important to get around the current strong self-pollination aspect of the species if hybrid soybeans are to be developed.

Effect of bees on soybean production and cross pollination
 Just as historically there was originally skepticism and confusion concerning soybean honey production, this has also been the situation with the topics: (1) how much cross pollination do honey bees accomplish? and (2) do honeybees increase soybean yield?

Cross-pollination 
 Originally it was dogma that soybeans were nearly totally self-fertilized. In the words of C. M. Woodworth (1922) [27], "When the stigma is receptive, the anthers burst open covering the stigma with an abundance of pollen grains." This statement was based on two experiments, each using a different genetic marker where the two lines to be crossed were systematically and intimately intermingled. In both experiments the crossing rate was only 0.16%. Piper and Morse (1923) in their book ‘The Soybean'[21] make similar claims, "The flowers are completely self-fertile, as bagged or screened plants set pods and seeds as perfectly as those in the open." As an indication of this, they cite two studies, one in Virginia (1909) where ten varieties were tested and a similar study in India (1913) that gave results identical to what they had just described. Milum (1940), at the University of Illinois, set wire cages over soybean plants at different distances from honeybee hives and found, ".....there were just as many seeds per pod beneath the cages as on the plants outside the cages." He concluded that "Since soybeans are self-fertile.....there should be little, if any, nectar available to attract the bees for the service needed for seed formation". Jaycox[13] in a review of the early soybean crossing literature, cites (without references) a work of Dr. Hadley in the Department of Agronomy at the University of Illinois who had done crossing experiments to assess the potential for cross-pollination. In what was apparently an open field test7, there were no differences in the crossing percentages between flowers in the upper and lower portions of the plant and overall, there had been only 0.39% crossing. In caged experiments, the cages with bees produced a crossing rate of 0.69%, while in the cages without bees no hybrid seeds were produced. Where Hadley conducted uncaged experiments near honey bee colonies, he found crossing rates between 0.21 and 0.47%. Jaycox (again without references) also provides data from experiments by Dr. Richard Bernard of the USDA Regional Soybean Laboratory at the University of Illinois, who had done experiments with an unusual noncommercial variety of soybean known for its ability to cross. Bernard found the crossing rate in this noncommercial variety was 15.5% when it was caged with bees and standard soybean varieties as sources of pollen. He also found the crossing rate was 11.6% in open field conditions when honey bee colonies were close by, but this percentage slipped to 6.6% when the colonies were farther away. Jaycox ends his review with the statement, "Standard soybean varieties grown in the Midwest, such as Clark and Harosoy do not benefit from visitation by honey bees. This is probably true also of all other commercial varieties."
 As I reviewed the literature, I encountered a translation of a Rus-sian paper[11] which provided an interesting insight into how difficult it is for plant breeders to make soybean crosses. The author, V. A. Gordienko, was interested in developing an easy and quick method for making soybean crosses using bees. If these crosses are made by a plant breeder in the field, they sometimes must lie on the ground, and then work under magnification with thin needles and forceps in order to remove the small and delicate stamens before pollen is released. This delicate operation must be done without adversely affecting the pistil, which at this time is pretty much surrounded by and in close contact with the staminal sheath (see the April column). Under these circumstances, the rate of stamen removal and subsequent pollen transfer was claimed to be about 30 flowers in an eight-hour day and the success rate of hybrid seed formation was only about 0.2%. When Gordienko[11] performed caged studies similar to those cited above, he claimed a crossing rate of between 28.6 and 44.1% in cages with bees. These were unexpected results and he suggested the unexpectedly high cross-pollination rate had resulted from either rapid changes in temperature and/or humidity within the cage, which weakened the corolla and caused it to split open prematurely before the pollen was released. He also provided another explanation that is unintelligible to me, perhaps because of a clumsy English translation.
 More recently, Abrams et al.[1] conducted an experiment designed to study the comparative effectiveness of honey bees vs. alfalfa leaf cutter bees (Megachile pacifica, now M. rotundata) for cross pollination and increased soybean yield. Six fields of purple flowered soybeans were used. In each field, perpendicular transects that intersected in the fields' centers were laid out, and small plantings of white flowering cultivars were planted at varying distances from the intersections of the transects. A honey bee hive was placed at the intersection of the transects in three fields and a commercial alfalfa leafcutter bee board was placed in a similar location in the other three fields. At the ends of the fields white flowered plants were also planted among the purple flowered plants. These plots were sprayed with insecticides and were used as control plots. Purple flower color is dominant to white flower color so that purple flowered progeny of white flowering plants would be the result of cross pollination. The honey bees were observed foraging actively within their fields and the colonies faired well, doubling in size and storing sufficient honey for winter maintenance. The leafcutter bees faired poorly and did not work the soybeans, but were observed actively flying from the field. At maturity both the white and purple flowering plants were harvested in all plots. Bean yields of the purple flowering plants were determined and the seeds from the white flowering plants were planted in a greenhouse to determine the rate of cross pollination. Soybean yields of the purple flowered plants were not significantly increased by either bee species. The cross pollination in the honey bee fields ranged from 2.95% to 7.26% compared to only 1.15% in the insecticide treated control plots at the ends of the fields. There was no generally diminishing trend of cross-pollination at the different distances from the hives, but I felt the data might be interpreted as having two peaks. There was, however, not an abundance of data points. In addition, the cross pollination in the alfalfa leafcutter fields, which appeared to not be worked by the leafcutters, ranged from 1.61% to 7.74% compared to 6.59% in the insecticide treated control plots. These last facts led the researchers to speculate on the presence of a third, but unknown pollinator.

Increasing production
 Erickson, perhaps spurred on by his recognition that the soybean flower appears to be designed to accommodate insect pollinators, has almost doggedly investigated not only soybean honey production covered partly in the April column and continued above, but also the effect of bees on soybean yield. During a three-year study in southern Wisconsin (1971-1973)[5] he investigated this topic using several varieties of soybean in cages with and without bees and also cages "without insects" where the soybeans within the cages were treated with an insecticide. Plots without cages served as control plots. Seed yield differences varied between both cultivars and years and appeared to result from differential attractiveness based on whether or not the flowers opened and probably also on differences in available nectar sugar. In the 1971 study the cultivar ‘Chippewa 64' never opened (cleistogamous) and produced no significant yield differences and was apparently totally self-pollinated. The cultivar ‘Carsoy' caged with honey bees yielded 13.9% more soybeans than those caged without bees. The insecticide treated vs. the control open, untreated plots showed no significant differences, but the plants caged with bees produced 14.9% more soybeans than the caged plants treated with insecticides. Both the 1972 and 1973 studies utilized ‘Hark' a relatively chastogamous8 cultivar. The 1972 study produced no significant differences between plants caged with and without bees. In the 1973 study the plants caged with bees produced 16.3 % more beans than the plants caged without bees. The difference between the insecticide treated plots and the open, untreated plots were apparently not significant, but the plants caged with bees produced 11.6% more beans than the insecticide-treated plots. Statistical significance in the 1973 trials was dependent on the statistical test used. 
 Wisconsin is approaching the northern limit of soybean production. To see how similar experiments as described above would play out farther south, Erickson et al.[9] in 1975 performed both "caged with and without bees studies" as well as "distance from hives" studies in both Missouri and Arkansas. The "caged with and without bees" studies were performed near Bragg City, MO and Jonesboro, AR using "Pickett 71' and ‘Pickett', respectively, in those locations. In combined results of the two locations, the caged plants with bees produced 21.6% more beans than the caged plants without bees and there was a 20.4 % increase in the total number of pods filled. Interestingly, open field plots, intended to serve as controls produced better than the caged plots with bees, which the authors believed resulted from the cages having a detrimental effect on the plants. Because they had not seen similar results in their Wisconsin studies, they suggested that caged studies should not be used in southern production trials. Their "distance from hives" studies were carried out near Wardell, MO using the cultivar ‘Forrest' and near Blytheville, AR using the cultivar ‘Lee 68'. In these experiments there was not a steady decline in soybean production with distance from the hives. Instead, production generally declined and then went through a secondary peak at about 250 meters and then again declined. The fields were not large uniform areas, but had significant landmarks (field edges and roads) and the authors felt that their data was consistent with the known foraging behavior of honey bees, which have been shown to forage heavily near landmarks, and they seemed to feel that they had demonstrated that there had been a decline in soybean production correlated with distance from the hives.
Kettle and Taylor (1979) [15] working in northeast KS found the "highly attractive" cultivar ‘Forrest' with a mean nectar solids9 concentration of 39.5% produced significantly greater seed yields of approximately 20% under cages with bees than under cages without bees.
Sheppard et al. (1979)[25] from their studies, presumably in Illinois, concluded that there was no relationship between soybean yield and distance from hives. As I look at their data, while there were not many data points, the pattern seems somewhat reminiscent of the "possible double peak" pattern found by Erickson et al.[9] and Abrams et al.[1], and I wonder if we are missing something. The Sheppard et al. paper provided the interesting bit of information that the Italian bee breeds seemed to forage over greater distances than the Caucasian or Carniolan breeds, suggesting that Italian bee strains may be more likely to forage flora outside of the soybean fields that is more attractive than soybeans.
Koelling et al.[16] (1981) examined the potential of honey bees vs. alfalfa leaf cutter bees for making hybrid soybeans using a male-sterile ‘Williams' line10 and a male-fertile ‘Calland' line. In these experiments they used cages with and without bees (the two bee species segregated into different cages) and as controls they used caged plots without either bees species and also plots that were caged, but with the cage sides rolled up to 60 cm (about two feet). Only the beans from the male-sterile plants were harvested. No bees of either species were added to the field in which the experiment was performed. The researchers found no significant differences in soybean seed production between honey bees and alfalfa bees. They also found no significant differences between closed cages and open cages. They did, however, find significant increases in seeds/plant (39.6 vs. 8.5) and pods per plant (19.1 vs.1.7) in cages with bees vs. those without bees. There was no significant increase in seeds/pod, indicating that the increase came from the number of pods that were set. Unlike the Abrams et al. study[1], they found the alfalfa leaf cutting bees more suitable, or at least easier to deal with, than the honey bees.
 Sheppard et al. (1979) [25] seem to suggest that indeterminate growth11 plants used in their study might not produce as much nectar as determinate plants commonly grown farther south than the indeterminate types. As a result, indeterminate plants would be less attractive to bees than determinate types, and bees would not, therefore, be expected to produce the benefits that they sometimes seem to farther south. Erickson[8] clearly disagrees with this assessment, stating, "I have yet to discern differences in foraging by bees or yield responses resulting from bee pollination that can be explained based upon level of determinancy at flowering." Instead, his work that demonstrated that cleistogamy is sometimes related to cool temperatures suggests to me at least that higher honey and seed productions in southern climates may be the result of warmer temperatures.

Recommended number of colonies per acre of soybean
 If bees can increase soybean yields, it is not well reflected in the pollination recommendations for the crop. McGregor[18] states "There are no recommendations for the use of bees in pollination of soybeans." He adds that he reviewed the literature primarily because of the interest in the production of hybrid soybeans. Delaplane and Mayer [4] state, "Supplemental bees are rarely, if ever used for pollinating soybean in the field. This could change if production of hybrids becomes practical, in which case bees will be needed to transfer pollen between parent lines." Scott-Dupree et al. [24] make a recommendation of "0" hives per hectare, but they do provide an estimate of soybean production being 5% dependent on honey bees and 10% dependent on insects as a whole.
 Jaycox[12] (1970) claims to have heard numerous stories of soybean growers in Illinois and other places wanting bees near their plantings, but was never able to collect hard evidence that this was true. Erickson[ 8] (1984), however states "Regardless of opinions to the contrary, many soybean growers continue to encourage beekeepers to locate apiaries near their fields and report increased yields with bees present." Ayers and Harman[2] reported some of the respondents to their questionnaires indicated that there was some commercial pollination of soybeans within the area for which they were reporting. In some cases, however, it was unclear exactly what was meant by "commercial pollination".

Prospects for hybrid soybean production
 Soybeans have become a very important crop, and there has been much interest in the production of hybrid soybeans. Like many of the other topics associated with soybeans, there is disagreement about whether soybean hybrids will become a reality. There are those who think they will be developed very soon, and there are those who think that it will never happen. From my perspective, the self-fertile nature of soybeans, the way that this is enforced by the flowers of some cultivars not opening, and the effects of the weather on floral opening all seem to be large problems that need to be overcome if hybrid soybeans are to become a reality. On top of that there are the more usual problems of developing satisfactory male-sterile plants, restorer lines, creating lines that the pollinators will move freely between, etc. Hybrid soybean seed has been created, but the yields have been disappointing and/or the process has been too expensive to compete with nonhybrid soybean seed production.
 In 2003 a group of Chinese researchers claimed to have produced the first practical hybrid soybean cultivar[28]. These crosses were apparently done with an insect other than the honey bee. My discussions with a U.S. soybean breeder indicates that the reported Chinese hybrid production system is very expensive and not economically competitive and that the hybrids do not produce as well as the best inbred lines. If this is true, it looks to me as though in the area of hybrid soybeans, we haven't yet arrived.

References
1. Abrams, R. L, C. R. Edwards and T. Harris. 1978 Yields and cross-pollination of soybeans as affected by honeybees and alfalfa leafcutting bees. American Bee Journal 118:555-556, 558.  
2. Ayers, G. S. and J. R. Harman. 1992. Bee forage of North America and the potential for planting for bees. In: The Hive and the Honey Bee (J. M. Graham, Ed.) Dadant and Sons. Hamilton IL.
3. Davis, J. H. 1952. Soybeans for honey production. American Bee Journal 92:18-19.
4. Delaplane, K. S. and D. F. Mayer. 2000. Crop Pollination by Bees. CABI Publishing. New York.
5. Erickson, E. H, 1975. Effect of honeybees on yield of three soybean cultivars. Crop Science 15:84-86.
6. Erickson, E. H, 1975. Variability of floral characteristics influences honey bee visitation to soybean blossoms. Crop Science 15:767-771.
7. Erickson, E. H. 1975. Honey bees and soybeans. American Bee Journal115: 351-353, 373
8. Erickson, E. H. 1984. Soybean pollinaion and honey production-a research progress report. American Bee Journal 124:775-779.
9. Erickson, E. H. G. A. Berger, J. G. Shannon and J. M. Robins. 1978. Honey bee pollination increases soybean yields in the Mississippi delta region of Arkansas and Missouri. Journal of Economic Entomology 71:601-603.
10.Erickson, E. H. and J. M. Robins. 1979. Honey from Soybeans: The influence of soil conditions. American Bee Journal 119:444-445, 448-450.
11.Gordienko, V. A. 1977. Obtaining sexual hybrids of soybean by controlled bee pollinatikon. In: Pollination of Agricultural Crops by Bees.[Mel'nichenko, A. N (Ed.). Vol 3 pp381-388. Amerind Publishing Co. New Delhi.
12.Jaycox, E. R. 1970a. Ecological relationships between honey bees and soybeans. American Bee Journal 110:306-307.
13.Jaycox, E. R. 1970b. Ecological relationships between honey bees and soybeans. II The plant factors. American Bee Journal 110:343-345.
14.Johnson, A. P. 1944. Honey from soybeans. American Bee Journal 84:306.
15.Kettle and Taylor 1979. Ecological interactions of honeybees and soybeans. J. Kansas Ent. Soc. 52:549 (abstract) 
16.Koelling, P. D., W. J. Kenworthy and D. M. Caron. 1981.  Pollination of male sterile soybeans in caged plots. Crop Science 21:559-561.
17.Lovell, H. 1957. Let's talk about honey plants. Gleanings in Bee Culture 85:228, 249,253.
18.McGregor, S. E. 1976. Insect Pollination of Cultivated Crop Plants. Agricultural Handbook Nol 496. Agricultural Research Service. United States Department of Agriculture. Washington D.C. This publication is being updated and is available on the web at: gears.tucson.ars.ag.gov/book/
19.Milum, V. G. 1952. Anet Soybean Honey. Report State Apiarist, Iowa pp. 53-55. Des Moines, IA.
20.Pellett, F. C. 1976. American Honey Plants, Together With Those Which are of Special Value to the Beekeeper as Sources of Pollen. Dadant & Sons. Hamilton, IL.
21.PiperC. V. And W. J. Morse. 1923. The Soybean. McGraw-Hill Book Co. Inc. Newyork.
22.Roebacker, D. C. , P. K. Flottum, D. Sammataro and E. H. Erickson. 1982. Why soybeans attract honeybees. American Bee Journal. 122:481-484, 518-519.
23.Robacker, D. C. et al. 1982. The role of flower aroma in soybean pollination energetics. Proceedings of the 10th Pollination Conference July 1982 Carbondale, IL. .pp 1-8.
24.Scott-Dupree, C., M. Winston, G. Hergert, S. C. Jay, D. Nelson, J. Gates, B. Termeer and G. Otis (EDS). 1995. A Guide to Managing Bees for Crop Pollination. Canadian Association of Professional Apiculturalists.
25.Sheppard, W. S., E. R. Jaycox and S. G. Parise. 1979. Selection and management of honey bees for pollination of soybeans. Proceedings of the 4th International Symposium on Pollination pp123-130.
26.Weiss, E. A. 2000. Oilseed Crops (second Edition) Blackwell Science, Inc. Malden, MA.
27.Woodworth, C. M. 1922.  The extent of natural cross pollination in soybeans. Journal American Society of Agronomy 14:278-283.

The Other Side of Beekeeping - April 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Soybean, A Good Honey Plant--Sometimes

Soybean, soya

Scientific name: Glycine max

Synonyms: Dolichos soja, Glycine gracilis, Glycine hispida, Glycine soja, Glycine ussuriensis, Phaseolus max, Soja hispida, Soja max

Origin: Glycine max is known only from cultivation (a cultigen), but is apparently closely related to the wild Glycine soja. The genus Glycine has two major gene centers, one in eastern Africa and the other in Australasia1, with a secondary center in China. While the origin of soybean is in dispute, it is generally thought to have originated in northeastern China.[8 &32]. Some believe that it became domesticated during the Shang dynasty (ca.1500-1100 BC) or perhaps earlier[8]. 

Plant description: Because the soybean is of considerable agricultural importance, the species now exhibits much variation due to the activities of plant breeders. In nature the soybean probably was much branched, but modern cultivars often have fewer than six branches.  The lower parts of the stem become woody with age.  The stipulate2 leaves are placed alternately on the stem and consist of three leaflets (trifoliate) that range in shape from broad and rounded (ovate) to longer, narrow and more pointed (lanceolate). Varieties with wavy leaflet edges (sinute) also exist.The leaf stem (petiole) is relatively long. Leaves are commonly dark green, but can be tinged with brown, red, or blue and are normally shed as the seed pods ripen. 
 Flowers are borne on short racemes3 in the upper angle between the leaf petiole and the stem to which it is attached (axil).  Floral groupings can contain up to 35 small typical pea-shaped white to mauve colored flowers. Frequently many of the flowers borne on very floriferous varieties abort without setting pods. The standard petal or banner petal is usually about 0.2 inches (ca. 5 mm) long.  There are two narrow wing petals and two tightly clasped together, but not fused keel petals that are shorter than the wings. The sexual column consists of the pistil and nine fused stamens (staminal sheath) and one single dorsal stamen. The nectary surrounds the ovary that usually contains 3-5 ovules and is in turn surrounded by the staminal sheath. The standard petal of mauve colored flowers have distinctive, more deeply purple-colored nectar guides that converge just above a tongue channel, which is located at the base of the standard petal and leads to the nectary. Within the tongue channel there are two tongue guides, one on either side of the single stamen, that guide the pollinator's tongue into the nectary area. While the white flowered plants have no purple nectar guides, Erickson and Garment[6] indicate that there are ultraviolet light reflecting patterns on both the white and mauve colored standard petals that probably also serve as nectar guides. In addition, the standard petal reflects ultraviolet light while the wing petals strongly absorb these wavelengths, and the two patterns together provide a strong contrast around the entrance to the tongue channel, and this may also serve as a kind of nectar guide.
 At times the stamen filaments elongate, so that when the flowers open, the stamens are nearly as long as the pistil when the anthers begin to release their pollen. Sometimes the elongation is sufficient to push pollen from the end of the keel. At other times, the flowers don't open and self pollination occurs within the closed flower. The number of flowers within a floral grouping that open simultaneously depends on the soybean cultivar. An individual flower remains open for only a single day[6, 7, 17 & 32]. 

Distribution: Soybeans are grown mainly in the North Central States with a "tail of production" that follows the Mississippi River south to about mid Louisiana (see reference no. 33). As of 2007, the states with the largest productions, in decreasing order, were IA, IL, MN, IN, OH, NE, SD, ND, AR[34]. Like many agricultural plants, soybeans generally don't persist in the wild[16] and would, therefore, not be expected to be found outside of cultivation in stands sufficiently large to be important in honey production.

Blooming period: Soybeans bloom in response to day length and temperature. The varieties grown in the US are divided into 13 maturity groups, each adapted to a narrow band between two latitudes that are only about 100 to 150 miles apart. The earliest maturity groups are adapted to northern Minnesota and southern Canada while the latest are adapted to southern Texas. The early varieties bloom when the days are relatively long and the nights are relatively short, whereas the later maturing groups bloom under relatively shorter days and longer nights.  Planting a variety further north than the latitude to which it is adapted will extend the period of vegetative growth and delay flowering and the opposite occurs when a variety is planted further south than the latitudinal range to which it is adapted4[35 & 36]. The beekeeping literature seems to suggest that soybeans grown outside of the latitude range for which they were developed are likely to be poorer honey producers than when grown within that range.
 Ayers and Harman[1] report blooming dates that ranged from June to October. For the IL, IN, IA and MO area a respondent to the Ayers and Harman questionnaires indicated the blooming period there ranges from mid June to the end of July. Another respondent from AR indicated the blooming period in that state extended from late June to near the end of September, the range being so long because there were so many varieties grown within the state.

The Other Side of Beekeeping - March 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Some More Members of the Asteraceae

Pearly-everlasting, western pearly everlasting, everlasting, life everlasting, straw-flower, moonshine, ladies' tobacco, silver-button, immortelle, anaphalide nacrée

Scientific name: Anaphalis margaritacea

Synonyms: Anaphalis occidentalis, Gnaphalium margaritaceum

Origin: The species is native to at least North America, but two references[6 & 16] indicate that Asia, probably northeastern Asia, is also part of the origin. Notice that its current distribution includes Alaska, suggesting that this probably is correct.

Plant description: Anaphalis margaritacea is a rhizomatous1 perennial that grows to heights of 12 to 36 inches. The pointy 1.2 to 4 inch long, stemless (sessile) leaves range from long and narrow with nearly parallel sides (linear) to less frequently a more spearhead shape (lanceolate). They frequently are greenish on their top surface and white and wooly (tomentose) beneath when young, often turning a rusty color with age. Both the whole leaf and higher resolution photos of the upper and lower surfaces are shown in the page margin. The two parallel black lines running through the leaves indicate the location of the high-resolution pictures. The species for the most part has its male and female flowers on different plants (dioecious) or there may be a few male flowers centrally located in a clump of female flowers. The involucre2 is about a quarter of an inch in diameter and the phyllaries are pearly white. You might at first mistakenly think the phyllaries as petals. For a better understanding of the structure of a flower in the Asteraceae see this column August 2005. The fruits are 0.5 to 1 mm achenes.3 The species is quite variable and the above is only a very general description. [4, 6, &16]

The Other Side of Beekeeping - February 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Safflower, false saffron, bastard saffron, Mexican saffron, carthamé des teinturiers, safran bâstard

Scientific name: Carthamus tinctorius

Origin: Probably Eurasia from the eastern Mediterranean to the Persian Gulf [11, 13 & 21]. The plant is known only from cultivation (a cultigen) [13 & 21].

Plant description: Safflower is a 2-6 foot high, glabrous1, shiny green annual with a vertical stem that branches in its upper parts. The leaves are sessile2, broadest beyond their midpoint, and have minutely spine-tipped teeth that make contact with the plant unpleasant. The involucre3 is 0.79-1.6 inch (2-4 cm) in diameter. The 15 to 150 flower heads of a plant are generally yellow to orange in color (rarely white or red) and range in diameter from 0.5 to 1.5 inches (1.3-3.8 cm) and occur at the upper end of the central stem (main flowers), at the ends of the branches (primary flowers) as well as along the branches (secondary flowers). Corollas4 of the individual florets are 0.78-1.18 inches (2-3 cm) long and terminate with five, pointed segments (corolla lobes) that together look a bit like a star. The highest flowers of the central stem open first with those on the branches opening progressively downward. Each flower head usually contains 20 to 100 florets where the outer florets open first, followed progressively by those in more inner positions. Flowering occurs over a period of 10-40 days with each flower head blooming for a period of 3 to 5 days. Nectar is secreted at the base of the stamen filaments.[8, 11, 13 & 15]

Distribution: Keil and Turner[11], writing about California plants, describe the plant's distribution outside of agricultural fields as disturbed places and roadsides primarily in the Great Central Valley and surrounding areas at elevations less than 1000m (3281 ft). When it escapes from cultivation, it occurs primarily as a waif5[13 & 21].

Importance as a honey plant: Ayers and Harman[1], from their questionnaires, found the species to be of some importance in CA and AZ. Howes[10] states, the flowers "secrete nectar very freely and are much visited by bees."

Honey potential: Eckert[7] reports honey yields of 30-60 lbs per colony by California beekeepers. Harvey Lovell[14] states that California beekeepers describe average per colony honey yields of 30 lbs. Boch[3] found the nectar sugar concentration to be 13-17% between 6.00 and 8.00 h, but became 24-29% the remainder of the day.
Pellett[16] provides the following quote from a personal letter from G. H. Vansell of the California Experiment Station at Davis, CA:
 
"A plot of it (safflower) grown here at Davis by the agronomy division attracts a greater number of bees in comparison to any other plant available at the present time. It produces an abundance of both nectar and pollen. An individual floret in the compound head produces so much nectar that it fills up the tube and runs out onto the bases of the petals. At times there are as many as eight bees per square yard estimated to be visiting this plot, which simply hums with activity. The nectar is quite rich in sugar, exceeding by 10 to 15% samples taken from neighboring alfalfa fields."

Honey: There seems to be some disagreement concerning the quality of safflower honey. Harvey Lovell[14] describes the honey as "high grade", rather dark but with a good to excellent flavor. Eckert[7] reports that the honey is "rather dark and strongly flavored" and when grown in the vicinity of alfalfa, produces a mixture inferior to that of alfalfa. R. B. Wilson[22], in Eva Crane's book ‘Honey a Comprehensive Survey', considers safflower honey from Arizona and California to be a one of the U. S.'s "miserable honey(s)". Elsewhere in the same book Crane states that safflower honey is "dark, strong (with an) unpleasant flavor and aroma" [5].

Pollen: Both the nectar and the pollen are highly attractive to bees[15]. Eckert[7] states that the pollen "appears to have an excellent brood-producing potential".

Additional information:
A Brief History
Safflower has been cultivated for many years and was being grown in Egypt at least as early as 2000 BC, but is a relative latecomer to mechanized arable cropping. The species was first cultivated for the production of two dyes, one yellow and one red. It was also recognized as having value as a potherb and for the production of oil used for cooking and medicinal purposes. It became an important oilseed plant in the US after World War II[21]. As an interesting aside to the dye story, one of the major uses of the red dye was to color cotton tapes that were used to tie legal documents together, and is suggested as the origin of the phrase "red tape"[23]. Both dyes have now largely been replaced by more stable synthetic dyes produced by the commercial dye industry. Today the species is primarily grown for its oil production. The yellow and reddish florets are sometimes dried separately to yield golden yellow and red powders that are used as a substitutes for the much more costly true saffron6 to flavor and color a variety of foods ranging from fish and seafood to salads and pastries[23].
Safflower is commonly cultivated in the old world and to some extent in North America, primarily in California and Arizona, but has been successfully grown in every state west of the 100th meridian (a line from mid-ND to mid-TX) [15]. Since 1975 the world production of safflower has been declining, being replaced by other oilseed crops (soybean, sunflower and the canolas)[21]. The species is, however, tolerant to drought and high salinity soils. These characteristics may help safflower become a more attractive alternative in the future because these environmental conditions appear to becoming increasingly more common[21].
Bee Activity and Pollination Requirements for Seed Production
In a Canadian study by Boch[3], bees commenced foraging at 07.00 h, becoming most numerous between 09.00 and 11.00 h, during which time estimates of nectar and pollen availability indicated that on-hand nectar and pollen supply decreased rapidly leaving only current production levels of both, and after 12.00 h the bee population decreased rapidly. This bee foraging pattern seems to be fairly general and was also observed by other researchers[12 &17]. In the Boch study, while the bee populations decreased rapidly after 12.00 h, the nectar sugar concentration did not. Data collected by Levin and Butler[12] suggested that the greater bee populations in the morning were independent of the size of the bee population. They found this surprising since lower bee populations should leave higher amounts of nectar and pollen in the field in the afternoon than would larger bee populations. These two observations taken together seem to suggest that something besides nectar concentration is determining bee populations in the safflower fields. Boch[3] hypothesized that the phenomenon was due to an increase in relative attractiveness of the surrounding environment (more competing bee forage), while Levin and Butler conjectured that there was some unknown attractive component of safflower that diminished in the afternoon. Whatever the cause, the generalized foraging pattern suggests that damage from pesticide applications would be less in the late afternoon than in the morning[12].
Levin and Butler[12] noted that both honey bee and other potential pollinator populations were considerably greater on the edges of safflower fields than in the middle, suggesting that if bees are used for pollination, they should be distributed throughout the field rather than along the field's edge.
In the Levin and Butler study[12] cited above, there were more nectar-collectors than pollen collectors, and while this may be the general situation, I suspect the relative number of foragers collecting nectar versus pollen is usually determined largely by conditions back in the hives. Rubis et al.[17] investigated an interesting example where the relative number of nectar and pollen gatherers apparently was not determined by hive conditions. These researchers investigated differences between two lines of safflower, one that produced a normal thick-hulled seed and the other a mutant thin-hulled variety that released its pollen a few hours later than the normal line. They found that pollen collectors worked only the normal line, but nectar collectors worked both. A few bees with pollen were seen working the thin-hulled variety, but they were thought to be basically collecting nectar. This difference in pollen release is the basis for an interesting and somewhat unusual method of hybrid seed production discussed in greater detail under ‘Hybrid seed production' below.
Safflower is generally considered to be a self-pollinated crop. Claassen[4], however, found that natural crossing of individual plants ranged from 0 to 100%. It is, therefore, not surprising that both McGregor[15] and Free[8] in their reviews of the literature, found evidence that the benefit of providing bees for pollination varied greatly. McGregor sites a two colony/acre recommendation made by Eckert in 1959[6] and then goes on to say that few beekeepers require payment for their pollination services of this crop because it is such a good nectar and pollen producer. He concludes, however, that only rarely are bees placed in safflower fields at densities as high as the Eckert recommendation, but that the grower would probably benefit more than the beekeeper by following the 2 colony/acre recommendation. Free concluded that bees would not greatly increase seed production in lines that are both self-fertile and self-pollinating, but where the line is missing either one of these attributes, bees would likely greatly benefit seed production. This is essentially what Eckert[7] in California and Rubis et al.[17] in Arizona found in their early safflower pollination studies.

Hybrid Seed Production
Historically hybrid safflower seed has been produced by an interesting method that pushes the pollination abilities of our bees to their limits. Initially the style and stigma are enclosed in a "tube" formed by 5 fused anthers that are attached to the corolla by short filaments (see accompanying diagram). Usually the style begins to elongate the morning the floret opens and pushes the stigma upward and out of the anther tube. If the anthers release their pollen before the stigma is pushed through the end of the anther tube, the stigma is coated with pollen as it emerges and self-fertilization can occur if the plant is self-fertile. If the pollen is released after the stigma emerges, there must be a transfer of pollen either from the anthers of the same flower or from another flower. Usually this is accomplished by some type of bee or more rarely by some other insect, as for example syrphid flies7. When the stigma emerges before the pollen is released, the flower is to some extent functionally male sterile (‘female') even though it is technically self-fertile. It is this situation that provides the interesting approach to the production of hybrid safflower seed alluded to above. A recessive gene dubbed ‘thin-hull' or ‘th' gene because it produces thin-hulled seed has been discovered that also provides this delayed pollen availability trait8. Once the pollen of the thin-hulled, ‘female' plant has matured, its anthers become fragile and are easily ruptured by bees foraging for nectar. For the thin-hulled plant to produce pure hybrid seed, it needs to receive pollen from a compatible plant before nectar foraging bees rupture its anthers and transfer its own pollen to the stigma. This window of opportunity is not very long (see Table 1), and to accomplish this pollination before the anthers are ruptured requires large numbers of foragers early in the morning.
As you might suspect, given the intricacies of this system, it doesn't work perfectly for hybrid seed production. Urie and Zimmer[19] found that pure hybrid seed produced in the greenhouse using hand pollination and pollen from the best variety of that time (Ute), outyielded that variety by 15 to 33% (average=24%). In the field, however, the crossing system described above does not normally provide pure hybrid seed, but also produces some of the thin-hulled, thth seed, which lessens this theoretical 24% figure (see associated figure). This problem isn't easily solved. In order to maintain the thin-hulled variety, which has the double dose of the th gene, there needs to be some self-fertility in that line, and as a result, some thin-hulled, nonhybrid seed is produced when the hybridization is carried out under normal field conditions. Remember, the window of opportunity for hybridization is not very long (Table 1). Apparently it was originally thought that the progeny of this thin-hulled contamination would be crowded out by the more vigorous hybrids, effectively making the outcome of the hybridization process pure hybrid seed. In the Urie and Zimmer study this didn't happen although the progeny derived purely from the ‘female' plant were overgrown. Using five normal lines as males and two thin-hulled lines as ‘females', Urie and Zimmer found that hybrid seed formed under caged conditions with bee pollination had 16 to 43% contamination by ‘female' selfs and sibs9. The progeny of these crosses were then planted under field conditions at five locations to study resulting yields. Using all the yield data, the average yields were 93.3% of the variety Ute. When only the results of when Ute was used as the "male" parent were considered, the yields were only 91.5% that of Ute. In addition, when the researchers planted normal hulled seed mixed with 10 to 60% thin-hulled seed (corresponding to the ‘female' of a hybrid crosses) they found that yields steadily decreased with increasing amounts of the thin-hulled variety. Even mixtures of only10% thin-hulled seed failed to yield as much as the pure stands of the two normal varieties used in the study. The result is that the hybrid advantage was essentially lost and unless the competition between the hybrids and the contaminating low yielding thth plants can be eliminated or at least reduced, the full potential of the hybrids will be lost.
Other approaches to the formation of safflower hybrids have been tried. Heaton and Knowles[9] in 1982 introduced a recessive gene (ms for male sterility) that resides in the nucleus. In its heterozygotic state (in combination with the dominant MS gene) the effect of the ms gene is totally masked, but in its homozygous state (msms) it produces no fertile pollen. Two germplasm releases of the ms gene were made available to other plant breeders.
In another approach, Baydar and Gökmen[2] found that three successive treatments of gibberellic acid (GA3) reduced pollen viability to as low as 6.7%. In their studies, hybrid seed production was 72.6% in main heads, 82% in primary heads and 87.5% in secondary heads10 with an overall average of 80.7%. It seems to me that unless the percentage yield of hybrid seed can be improved, the resulting progeny might suffer from the same problems uncovered by Urie and Zimmer[19]. In addition, gibberellic acid is a plant hormone that has many effects on treated plants and the treatments used by Baydar and Gökmen apparently resulted in a higher hull percentage and lower oil content of seeds from treated plants than from nontreated plants. These authors also suggest that there may be effects of the treatment that show up during the germination of the hybrid seed.
To date, there seems to be no perfect solution to the creation of safflower hybrids. Weiss[21] summarize the situation as, "Cytoplasmic male sterility would greatly assist breeders as it has done with other oilseed crops."11
As is always the case, no matter which hybrid seed production system is utilized, steps need to be taken so that foragers do not become conditioned to (prefer to work) one of the two lines. Rubis [18] found that bees could become conditioned to lines based on floral color. Also, when the numbers of ‘female' to male rows were planted in ratios of 2:2, 4:2, 8:2 and 18:2 he found that the percentages of cross-pollination were 71, 63, 52, and 32 percent respectively, and in the 18:2 blocks the percentage of cross pollination decreased toward the central ‘female' rows but was 79% in the ‘female' rows that were adjacent to the two blocks of male rows, suggesting that the male and ‘female' lines should be planted in alternate rows.
I personally wonder if interest in production of safflower hybrids will abate given the present decline in safflower production and the spectacular successes that are occurring in competing oilseed crops.

References 
1.Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In the Hive and the Honey Bee (J. M. Graham, Ed.) Dadant and Sons. Hamilton IL.
2.Baydar, H. and O. Y. Gökmen 2003.  Hybrid seed production in safflower (Carthamus tinctorius) following the induction of male sterility by gibberellic acid. Plant Breeding 123:459-461.
3.Boch, R. 1961.  Honeybee activity on safflower (Carthamus tinctorius L.) Canadian. Journal of Plant Science 41:559-562.
4.Claassen, C. E. 1950.  Natural and Controlled Crossing in Safflower. Carthamnus tinctorius L. Agronomy Journal 42:381-384.
5.Crane, E. 1975. The flowers honey comes from. In Honey A Comprehen Survey (E. Crane Ed.) Crame Russak amd Company, Inc. NY.
6.Eckert, J. E. 1959. Honeybees in Crop Pollination. California Agricultural Experiment Station Service Leaflet 32, Revised.
7.Eckert, J. E. 1962. The Relation of Honey Bees to Safflower. American Bee Journal 102:349-350.
8.Free, J. B. 1993. Insect Pollination of Crops (Second Edition).  Academic Press Inc. San Diego, CA.
9.Heaton , T. C. and P. F Knowles. 1982.  Inheritance of Male Sterility in Safflower. Crop Science 22:520-522.
10.Howes, F. N. 1979. Plants and Beekeeping.  Faber and Faber. London.
11.Keil, D. J. and C. E. Turner. 1993. Carthamus Distaff thistle.  In: Hickman, J. C. (Ed.) The Jepson Manual. University of California Press. Berkeley, CA.
12.Levin, M. D. and G. D. Butler Jr. 1966. Bees associated with safflower in south central Arizona. Journal of Economic Entomology 59:654-657.
13.Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dictionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co.,Inc. New York.
14.Lovell, H. B. 1966. Honey Plants Manual: A Practical Field Handbook for Identifying Honey Flora. A. I. Root Co. Medina, OH.
15.McGregor, S. E. 1979. Insect Pollination of Cultivated Crop Plants.  Agricultural Handbook 496, Agricultural Research Service. United States Department of Agriculture. Washington DC.  This publication is being updated and is available on the wet at: gears.tucson.ars.ag.gov/book.
16.Pellett, F. C. 1978. American Honey Plants. Dadant and Sons, Hamilton, IL.
17.Rubis, D. D. , M. D. Levin and S. E. McGregor 1966. Effects of honey bee activity and cages on attributes of thin-hull and normal safflower lines.  Crop Science 6: 11-14. 
18.Rubis, D. D. 1970. Bee-pollination in the production of hybrid safflower.  Report of the 9th Pollination Conference 43-49.  Hot Springs AK.  University of Arkansas and USDA.
19.Urie, A. L. and D. E. Zimmer.  1970. Yield Reduction in Safflower Hybrids caused by Female Selfs. Crop Science 10:419-422.
20.USDA, NRCS. The Plants Database. Verwion 3.5 (http://plants.usda.gov). National Plants Data Center, Baton Rouge, LA. 70874-4490 USA
21.Weiss, E. A. 2000. Oilseed Crops. Blackwell Science Inc. Malden, MA.
22.Wilson, R. B. 1975. World Trading in Honey.  In Honey: A Comprehensive Survey (E. Crane Ed.) Crame Russak amd Company, Inc. NY.
23.W. J. Beal Botanical Garden. Interpretive signage that accompanies most of the more than 2000 taxa within the garden. The Gardens are located on the Michigan State University Campus in East Lansing, MI.

The Other Side of Beekeeping - January 2010

Excerpt

Family Lythraceae--the Loosetrife family

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

The Lythraceae consists of herbs, shrubs, and trees, containing about 25 or 26 genera, and depending on the reference, somewhere between 550 and 580 species. While the family is widely distributed, it is largely tropical, but its members can be found in all but the very coldest parts of the world, and there are seven genera native to the U.S.
 The leaves are generally attached oppositely to their branches, but sometimes alternately in the upper parts of the plant. Frequently stipules are lacking, and when they exist, are small. The branches to which the leaves are attached are frequently four-sided.
 The flowers can be solitary, in that case are found in the upper angles between paired leaves and stems (upper axils), or they can be arranged in various types of floral clusters (racemes, panicles or cymes).
 The individual flowers are bisexual and generally at least roughly radially symmetrical, but occasionally might be considered bilaterally symmetrical. They are strongly perigynous, that is, they have a cup-like structure (hypanthium) around the ovary that appears to be formed by the lower parts of the calyx, petals and stamens (see diagram). They generally have 4 or 6 sepals (sometimes 8) and either an equal number of petals or no petals. Where petals occur, they are often crepe-like. In this family, while the petals and sepals appear to arise from the upper edge of the hypanthium, the stamens arise from deeper within the structure1. Generally there are twice as many stamens as sepals, but occasionally there can be many or as few as one or two.
 The stamens are frequently quite variable in length, often in the same flower. The pistil is compound, made up of 2 to 6 carpels2 with a single style and a stigma that usually is shaped like the head of a pin (capitate), but occasionally is two-lobed. It is sometimes buried deep in the hypanthium. Like the stamens the stiles vary considerably in length between plants, and according to one author[4], even on the same plant. The ovary is placed above the base of the hypanthium (superior). The fruits are capsules3.
 Some members of the family make good garden and greenhouse ornamentals. In the past some have found medical applications, some have been used in the perfume industry, and some in the dye industry. In the tropics, members of the genus Lagerstroemia are used for timber. Other members of this genus, grown in more temperate regions, make exceptional ornamental shrubs or small trees (example crape-myrtle). [4, 8, 9 & 24].

The Other Side of Beekeeping - December 2009

 (excerpt)

Rape/Canola: Two Very Productive But
Potentially Problematic Bee Forages    Part II

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Honey: Crane et al.[3] say that B. napus honey is bright yellow, pale yellow or golden.. They report the Pfund value as water white, 35mm white1, and light amber when liquid. They also provide the warning that it granulates very quickly-within a few days and sometimes even in the comb, The consistency of the crystallized product is fine and homogeneous, the color then white. The flavor of the ungranulated honey is reported as sweet and mild. When granulated, it is described as having little flavor and also as delicate. The aroma is said to be like that of the flowers or that it varies from none to rather unpleasant. The following sugar composition is reported:
glucose: 35.13 to 42.6%
fructose: 36.9-40.2 %
sucrose: 0.40%
maltose: 5.80 %
 Larson and Shuel[15] describe the honey as being "very white", having a mild flavor, and having a very high glucose content, causing it to granulate rapidly.
 Palmer[23], using the names Argentine rape and black Argentine rape, suggesting B. napus, cautions the readers about the honey's rapid granulation, but then states, "When granulated it contains no granules, like most honeys, but is as smooth and with much the same texture as butter. It spreads like soft butter all winter. It is similar to creamed honey, made with the Dyce process, the only difference being that rape honey granulated naturally that way." In 1959 he or someone with the same name and living in the same Saskatchewan town, and indicating that he had been producing rape honey since 1944, quotes some of his customers as, "the loveliest white honey I've seen", "so easy to spread" and "can't buy anything as good in stores"[24]. The 1959 paper claimed good overwintering on the honey, claiming successes in the winter of 1958-59 when there were no flight days for 112 consecutive days, and in the winter of 1955-56 when bees had been confined for 142 days, when in his words, there "was no loss or weak hives".
 White et al.[30] provide a set of honey characteristics under the synonym B. campestris, (a synonym of B. rapa), but the fact that there was no granulation after 17 months leads me to question whether this single sample was, in fact, from B. rapa.
 Crane et al.[3] under B. campestris, states that the honey color is white or yellowish white, clear or yellowish. They seem to question whether the granulation is rapid (their expression: "?rapid"). Perhaps this question stems from the granulation properties supplied by White et al. They report the flavor as "like wine", and "rather like the plant".
 R. S. Walsh[29], an Apiary Instructor in the Christchurch region of New Zealand, while not providing a species identification, indicates that the color of the honey appears to vary with soil type. The honey from South Canterbury is quite uniform, but farther north, where the land is heavier, variation in color is reported. He also describes how the honey granulates after extraction in about seven days and with even slight agitation, will crystallize in only four days. Frames of honey not removed from the hive will granulate within the comb in about a month. The honey after granulation he reports as water white in color with a fine grain and mild flavor.
  After reviewing the literature, my conclusion is that the honeys of B. napus and B. rapa are most likely quite similar.
 The rapid crystallization of canola honey is its major problem. If you are the rare producer who markets most of your honey as a ‘creamed' product, its rapid crystallization will not present much of a problem, it may even be desirable. On the other hand, if you are a typical U. S. beekeeper having your bees mix canola honey with one of your high quality honeys, say a clover honey for example, this may present a serious problem. It apparently doesn't take much canola honey, perhaps as little as 10 to 20%, to cause a whole batch of otherwise high quality liquid honey to granulate. This, I suspect, would be particularly true if a small amount of canola honey that is beginning to crystallize is topped off by the bees with another honey. As the amount of canola grown in the U.S. increases, as I expect it will, this rapid crystallization property will pose a problem until U.S. beekeepers learn how to manage canola honey much as the Canadian beekeeping industry apparently has.

The Other Side of Beekeeping - November 2009

Rape/Canola: Two Very Productive But
Potentially Problematic Bee Forages    Part I

(excerpt)

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Because discussions about the two species of rape covered here are often linked, and because it is often not clear about which plant is being discussed, I have linked them together in this writing using a format similar to the one I usually use. Where I can, I distinguish between the two species, and where I can't, I indicate this.

Rape, Argentine rape, rapeseed, oilseed rape, swede rape, summer rape, winter rape, colza, colza-oil rape, canola (only certain cultivars).

Scientific name: Brassica napus

Field mustard, summer turnip rape, Polish rape, toria, sarson, yellow sarson, colza, winter rape, canola (again only certain cultivars) navette

Scientific name: Brassica rapa

Synonyms: The term canola is a trademark of the Canola Council of Canada and represents the contraction of, Canadian and ‘ola', ola indicating oil. The name was originally applied to two species within the genus Brassica (B napus and B. rapa), whose seed met the standard of containing 2% or less erucic acid¹ in its oil and whose dry matter, after oil extraction, contained less than 30 micromoles²/gram of a mixture of glucosinolates³. This definition has been extended to include other plants within the genus Brassica, if and when their seeds meet these standards. It appears that Brassica juncea (India mustard) either has, or soon will meet these standards and part of that species will then also be referred to as canola. While the development of canola from rape⁴ has had a very large impact on the oilseed industry, it has, in my opinion, also led to considerable confusion taxonomically, since much of both of the two major species are not canolas since they do not meet the above definition. The situation is even more confusing because they both have annual and biennial forms often referred to as spring and winter rapes or canolas, respectively. The annual form is planted in the spring and harvested that year, while the winter form is planted in the fall and harvested the following year. In addition, there have historically been numerous synonyms applied to the two species, and B. juncea will add to this list. 

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The Other Side of Beekeeping - October 2009

The Brassicas 

(excerpt)

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Cabbage, broccoli, cauliflower, kohlrabi, kale, Brussels sprout
(all with various forms and names)

Scientific name: Brassica oleracea

Synonyms: To this day I remember when, as a young undergraduate, I was informed by one of my professors that cabbage, broccoli, cauliflower and Brussels sprout, etc. were all the same species. I was more than a little shocked! It wasn't, however, until recently as I began to think about this particular article, that the corollary to this bit of information, that genes can easily be passed between these entities, entered my mind and assumed any significance (see Additional Information below). Often discussions about B. oleracea divide the species into groups. In other writings the groups are treated as classical botanical varieties and sometimes even as species. A synopsis of these treatments is provided in Table 1.

Origin: While it is not always totally clear from where the various groups originated, on the whole they apparently came from Europe and Asia.

Plant description: In addition to the short descriptions provided in Table 1, cole crops are large-leaved plants that in their vegetative state usually grow to heights of about 1 to 2 ft until the flowering stem is formed, at which point they may grow to 7 ft in height with numerous branches and small leaves that come mainly from the main stem. The flowers on these stems are numerous, generally yellow but occasionally white, and have the four-petal arrangement and the "4 long, 2 short" stamen distribution, both characteristic of the family (See this Column September 2009 ). In most of this group, nectar is apparently secreted mainly by two nectaries located between the two short stamens and the ovary. There are also what have been described as two netaries located outside the two pairs of long stamens. It has been said that these are functionless_{[17 & 20]}, but Free and Williams_[10] describe bees "robbing" nectar from them (see Additional Information below), which suggests that they are not always functionless.

Distribution: Members of the species are usually grown in cool climates or during cooler parts of the year in warm climates, and do best under relatively high humidity_[16]. While many of the Brassica oleracea groups are grown throughout much of North America, my interpretation of the USDA Plants Website definition of "not in PFA¹" suggests to me that most do not, without human intervention, escape ivation and become permanently established (naturalized).

Blooming period: Generally cole crops are biennials that do not bloom until after they have gone through a cold period (vernalization). Usually they flower the year after they are planted and harvested. There is a fair amount of variation in this trait even within a group. Some varieties, especially those from warmer climates, when grown in temperate parts of the world, may assume an annual behavior. Most of us who have grown broccoli in our gardens, for example, have seen it bloom the year of its planting, especially after the main head has been removed.

Importance as a honey plant: Oertel[18], from his questionnaires, found cabbage to be important in WA, and kale to be important in OR. Ayers and Harman_[1], from their questionnaires, found cabbage to be important OR and WA, broccoli to be important in OR and AZ, and cauliflower to be important AZ, with all cases reporting at least some commercial pollination. Pellett_[21] has this to say about broccoli:

"In the Rio Grande Valley of Texas broccoli is extensively grown as a winter crop and blooms freely from the branches after the heads are cut. During the months of January to March the bees work broccoli freely and apparently harvest considerable honey from this source." Much of the commercial broccoli production now occurs in California and Arizona.

Concerning cabbage as a honey plant he says, "In the seed belt of California, where grown for seed on a large scale, cabbage is valuable".

More recently, Burgett et al._[2] writing about areas of commercial cabbage seed production in Oregon, state that it is, "An excellent honey plant with several thousands of acres devoted to commercial seed production."

McGregor_[16], about the species in general, states, "The flowers are highly attractive to pollinating insects for both nectar and pollen. When the seed-producing acreage is large, beekeepers nearby frequently harvest a crop of excellent honey."

Honey potential: Pearson_[20], apparently meaning to describe the nectar secretion for the B. oleracea groups in general, provides the information that it is about 0.1 ml each 24 hrs for the three days that the flowers are open.

Free_[8] from his review of the literature, provides the following estimates relative to honey potential:

For cabbage: 100 to 400 mg nectar; 0.5 to 1.0 mg nectar; and 3.1 to 4.9 mg/flower/24 hrs and a sugar concentration of 30-59% for 3 days, thus providing an estimated 12.5-15.1 lbs sugar per acre (14-17 kg sugar/ha).

For kohlrabi: 5.0-6.5 mg nectar/day with a sugar concentration of 30%, thus providing an estimated 33-46.3 lbs sugar/acre (37-52 kg sugar/ha).

Honey: McGregor_[16], apparently talking about the qualities of B. oleracea honey in general calls it "excellent". Burgett et al._[2], under the heading of cabbage, state that the honey granulates quickly. Pellett[21], referencing J. S. Harbison_[12], provides the following quote "Cabbage blossoms afford a considerable amount of honey of fine quality and flavor-Beekeeper's Directory." Interestingly Pammel and King_[19], apparently quoting the same reference by Harbison, state, "Cabbage blossoms afford a considerable amount of honey of fair quality and flavor."² When I decided to see for myself what Harbison actually said, Pellett got it right³. Harvey Lovell_[15] says that cabbage grown for seed in Washington produces a white, mild honey.

Pollen: The various groups of B. Oleracea provide pollen to our bees.

Additional Information: Generally B. oleracea crops are harvested before their flowers open. In fact, in many cases the plant must experience a cold period, referred to as vernalization, before it will produce flowers. In the field, this often means that it will not flower until the year following its planting and long after the crop is harvested and probably also turned under in preparation for the next crop. Beekeepers will, therefore, frequently be interested in providing pollination services for seed production rather than strictly for honey production.

Footnotes
¹ PFA: Plants Floristic Area
² The underlining in these two quotes is mine to draw attention to the difference between them.
³ Tracking down this reference turned out to be an interesting, even exciting, side venture. If you like reading about the history of beekeeping, the full text of this book can be found on the web (see references for its web address) Among other things, be sure to read Harbison's account of moving bees to California by boat during the Gold Rush days; and there is much more.

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