So the insect had little chance of defending herself when Johnson, a 31-year-old graduate student of entomology at the University of Illinois at Urbana-Champaign, plucked her from the cube and tossed her into a misty cooler filled with dry ice. Landing on the solid chunk of carbon dioxide, the bee gently folded her legs as she succumbed to the vaporous cold.
Working quickly, Johnson guillotines the insect with a firm pinch to the taper separating head from thorax. He peers into an aging gunmetal-gray microscope, then inserts the fine point of a pair of scissors at the base of the bee's abdomen. With a few clicks of thumb to forefinger, he gingerly cuts away the animal's thin abdominal casing, exposing the immature honeybee's ochre-colored viscera.
It will be the day's first dissection.
To the untrained eye, the string of organs Johnson lays bare looks more like snot than a discrete honey crop and intestinal tract. But Johnson, a baby-faced researcher who has spent the past few months scrutinizing the brown and runny innards of insects frozen as far back as 2005, rarely sees such a healthy specimen.
"This is so much easier on a fresh bee," marvels the entomologist, whose long-limbed and slender build could easily be mistaken for that of a teenager, were it not for his full head of prematurely gray hair. "Now I'll take this gut and refreeze it."
Toiling in the lab of professor May Berenbaum, Johnson is part of a nationwide effort launched by entomologists to solve the mysterious mass die-offs that have been plaguing the country's honeybee population. First reported this past November, the phenomenon known as Colony Collapse Disorder, or CCD, is marked by hives whose population of worker bees simply does not return home, leaving the queen and her immature brood to languish and die.
The mysterious disappearances have prompted entomologists from Pennsylvania to California to scour empty beehives and dissect the survivors in hopes of finding a culprit. Some teams are looking into whether pesticides are to blame. Others are searching for a new or emerging pathogen. Still more are examining honeybee nutrition and the host of stresses that mark modern apian life.
So far it's a bust.
The work has gained urgency because of the vast scale of the disappearances. The honeybee population is constantly growing and shrinking, making it difficult to put a precise number on the total population, but scientists estimate that CCD has felled roughly 25 percent of the nation's honeybees. The phenomenon has been reported in 37 states to date, as well as in Canada and Europe. The Apiary Inspectors of America, the official association of state regulators, reports that of the 384 beekeeping operations reporting losses, nearly a quarter lost more than 50 percent of their colonies over the winter. Some lost as much as 90 percent of their hives.
That's a lot of bees. It's also a lot of money. So far CCD has cost beekeepers an estimated $150 million. But beekeepers say that's just a foretaste of the devastation Colony Collapse could wreak on a pollination industry valued at roughly $14 billion.
Honeybees play a vital role in the agricultural industry. While they're not the only means by which crops are pollinated, they are uniquely suited to the task. Honeybees are easily transportable. Hives can be grown or shrunk to fit the demands of the job. Perhaps most important, honeybees are incomparably versatile capable of efficiently pollinating more than 100 different fruits and vegetables. What's more, they are the only pollinator capable of delivering in an agricultural system where acres upon acres of identical crops must be pollinated simultaneously.
But honeybees have paid a price. A non-native creature imported to the United States from Europe in the seventeenth century, they've lost their nature the feral honeybee population was wiped out more than a decade ago and exist today less as a species than an agricultural commodity, as essential as fertilizer.
And infinitely more mysterious.
"People have just assumed that pollination would always be there, like clean water or fresh air," says Berenbaum, who's also a member of the CCD working group. "The main finding so far is that we just don't know very much."
Modern American beekeeping bears only a passing resemblance to the once-gentlemanly pursuit of honey-making. Beekeepers continue to harvest honey as a natural and lucrative byproduct of keeping bees, but these days it's chiefly a sideline to their real business: pollination services.
An insect harvesting nectar from a blossom invariably rubs up against the flower's stamen. When the insect moves on, it takes some pollen with it. At the creature's next destination, some of the pollen from the prior encounter rubs off onto the blossom's female reproductive organs, or carpels. Though many insects serve as pollinators, a honeybee's anatomy, coupled with its industrious nature, make it ideally suited to the task. While the sex life of plants is protracted and nearly invisible, crop farmers will pay beekeepers handsomely for their efficient intervention.
So it is that early on a dew-kissed morning in southeastern Missouri's bootheel, beekeeper Neal Bergman is hunched over the flattened front tire of a forklift. The air is still cool from the previous evening's rains, and the 300 or so hives he keeps at his compound's central bee yard outside the town of Kennett have yet to rouse for the day. They will soon, and Bergman needs to move quickly.
Bergman's outfit, Delta Bee Company, owns several of the work-worn lifts, and by noon his 8-man crew will have moved 450 hives onto area watermelon patches. The faster they can do it, the less time the bees will have to awaken, get angry and sting.
"The way I look at it," says Bergman, a tall, broad-faced man whose voice retains the flattened vowels of his Minnesota heritage, "you're going into a houseful of women and you're rearranging the furniture. So when you're going in there, you're literally going into a hornet's nest."
There's another reason for Bergman's haste: Once the bees wake up, the workers will head into the field to do what Bergman employs them to do: forage for nectar and pollinate. Move the hives while the workers are away, and they won't be able to find the hive when they return with nectar. If left behind, the bees are likely to die of exposure.
Having changed the tire, Bergman straps himself into his Ford flatbed truck. One by one his crew departs, a circuitous single-file parade of five flatbeds through the region's patchwork of cotton and watermelon fields.
A few minutes later, the crew arrives at Delta Bee's main holding yard, where it is met by a bee-stung and swollen-teated Jack Russell terrier tiptoeing nervously through the moist morning grass.
It's at this central yard that the beekeeper consolidates his hives before shipping them out to fields whose owners pay him to pollinate everything from apples to zucchini. Bergman keeps roughly six hundred of the two-foot-by-two-foot wooden boxes at his central yard. Each hive is equipped with removable honeycomb frames, which allow Bergman control of the hives' strength, removing bees as necessary. The bee yard is studded with several 35-gallon plastic barrels brimming with high-fructose corn syrup used to "slop feed" the bees.
The yard's low-frequency buzz, ornamented by the morning song of cardinals, is quickly doused by the forklifts' roar. As Bergman's rig carves tread marks into the moist ground, his crew moves from hive to hive, perfuming the air with smokers, bellows-like canisters that belch smoke from smoldering pine needles to mollify the bees. The workers, dressed identically in white beekeeper suits, lend the operation a vaguely extraterrestrial quality.
Bergman, on the other hand, wears jeans and a button-down denim shirt with a Sue Bee Honey logo, a nod to the cooperative that buys most of his product. He has covered his head with a veil for protection, but otherwise Bergman leaves himself open to the elements as he expertly pilots the forklift, scooping the wood-sided hives six at a time and dropping them on the flatbed.
The son of Minnesota dairy farmers, Bergman now runs Missouri's largest beekeeping operation. He keeps an active stock of 8,000 hives, each of which can hold as many as 60,000 bees. He has a grain-silo-size tower filled with high-fructose corn syrup, which he buys by the tanker load as supplemental feed. This past February Bergman loaded 3,000 hives onto tractor-trailers bound for the California almond blossom.
"These bees are already world travelers. They've been to California for almonds, and they went up to Illinois for apple pollination," says Bergman. "This last year the [almond] rate was anywhere from $120 per hive to $160 per hive. That sounds great, but by the time you take out your broker fee for placing them, and for transportation it's about $10,000 per semi it gets pretty costly."
Expenses notwithstanding, the California almond blossom is by far the most important season for a migratory beekeeper like Bergman. When the almond trees, all 420,000 acres of them, begin to bloom in February, beekeepers from across the United States load their product onto flatbed tractor-trailers and ship the bees west. Each individual blossom must be pollinated, and collectively the almond growers need upward of one million bee colonies to accomplish the task. For beekeepers like Bergman, it represents the year's biggest payday. But it comes with its attendant perils.
"You put a hive of bees on a truck and run it 2,000 miles, you'll put that hive in extreme elements. They may experience anything from sub-freezing temperatures to temperatures over 100 degrees," says Bergman, who each January begins preparing his bees for the journey. "When we send the bees out there, we give them a shot of feed first and a protein supplement. Then we give them an external mite treatment. We give them an internal mite treatment. We give them something for nosema [a digestive problem], and we give them an antibiotic. We give that to them all at once, so when they head out there they have the whole drugstore thrown at them."
They need it. From a bee's perspective, pulling hives together from around the country is not unlike bringing together a million midsize cities and commingling their populations for a month. If one of those towns is experiencing a plague, there's bound to be some cross-infection.
But today's trip, like most that take place during the pollination season from February through May, won't involve interlopers. By 9 a.m. the air in the bee yard is thick with swarming bees and the smell of smoldering pine. But soon each flatbed is stacked high with hives. Bergman's crew lashes down the cargo with nylon webbing, and the trucks depart for watermelon fields as far south as the Arkansas border, and north into the region around Cape Girardeau.
As far back as the nineteenth century, beekeepers struggled with American foulbrood, a bacterial infection that kills bee larvae. Taking notice, Congress passed the Honeybee Act of 1922, which aimed to protect the nation's bee population from disease by strictly regulating the importation of foreign stock. But the Honeybee Act (which ultimately was repealed in 2004) failed to prevent several mysterious die-offs during the twentieth century.
Of course, bees have been susceptible to mysterious mass kills since their introduction to the United States in the 1600s. But although these die-offs, which went by names like Fall Dwindle Disease, Spring Dwindle Disease and Disappearing Disease, were recorded, beekeepers and scientists failed to keep detailed records of the symptoms or how many bees were lost. So today's researchers don't know whether CCD is worse than these earlier plagues, or even whether the phenomenon's symptoms are similar to any of the earlier epidemics.
"Nobody ever figured out what the earlier disappearances were. The records are very sketchy," says University of Illinois entomologist May Berenbaum, who last year authored a study on the state of the nation's pollinators. (Not good.) "We may be bad now, but at least we are making an effort to keep records."
The U.S. honeybee population took a severe nosedive in the mid-1980s. The cause was a one-two punch of nonnative mites: the varroa mite, which attacks bees during their larval stage and literally sucks the life out of them, and trachea mites, which live in the insects' respiratory tubes and interfere with breathing.
Intended to protect the domestic honeybee population, the Honeybee Act of 1922 had at least one unintended consequence: It effectively froze the U.S. honeybee population's genetic variability. There were so few new bees entering the country that by the time the varroa mite arrived the far more devastating of the aforementioned two pests the U.S. honeybee population was a veritable monoculture made up of hive after hive of genetically identical bees.
Worse, those bees had never encountered varroa mite, which some scientists estimate helped reduce the nation's domesticated honeybee population by 50 percent since it appeared on these shores in 1987.
Even a beekeeper like Bergman, who has not felt the effects of CCD, says that varroa mite has taken a heavy toll on his operation: Like many beekeepers, his annual mortality rate hovers at roughly 30 percent unthinkably high for most modes of agriculture.
"It used to be that you had, like, a 10 to 15 percent mortality rate," Bergman says. "Now with varroa and trachea mite and hive beetle and insecticides it just gets to be a lot of stress on bees."
If the mite has taken a heavy toll on the domesticated honeybee stock, it has destroyed the country's feral bee population. Bereft of manmade mite treatments, feral bees have almost entirely disappeared from the landscape, leaving today's post-varroa honeybee a creature almost wholly dependent upon man for its existence.
"The varroa was a huge hit. It removed this background of feral bees," says Charlie Whitfield, an associate professor of entomology at the University of Illinois. "The feral populations were not getting treated for mites. They'd had to deal with pesticides on their own, so there was a fair amount of natural selection going on. But once a new pathogen comes along and you don't have the variant present in your population to adapt, everybody dies."
Though unknown domestically until it was introduced to North America by Dutch settlers in 1638, apis mellifera has a long and intimate history with its human keepers. Cultivated by humans as far back as 4,000 B.C., honey is also responsible for mead likely the first alcoholic beverage. Egyptians used beeswax for embalming, mummification and sealing coffins, and several early societies used wax and honey the latter because of its antibacterial properties to dress wounds.
The advent of beeswax candles was an early invention that would carry over into Christian religious services and prompt the long history of beekeeping monks. More recently, European settlers were unable to cultivate apple trees until a group of seventeenth-century Dutch settlers imported honeybees.
In the years following their arrival in the United States, beekeepers imported several different varieties of honeybee from as far away as Japan and Russia. By the late nineteenth century, though, the so-called Italian Gold had become the bee of choice. Prized for its gentle disposition and prodigious honey-making ability, the Italian Gold strain is now predominant in the U.S. honeybee stock, although U.S. honeybees are so interbred at this point that it's a bit of an apian mutt.
Aside from the obvious benefits of beeswax and honey production, it is the social structure of the hive self-contained, with consistent, manipulable biological rhythms that has ensured the close relationship between humans and honeybees.
"Honeybees are the one species we can manipulate, and manage, and transport, package and deliver in a timely manner," says May Berenbaum. "They are exceptionally well suited to industrial agriculture."
Though pollination depends upon an individual bee flitting from plant to plant, leaving a fertilizing trail of pollen as it gathers nectar, a single honeybee in and of itself is not a viable organism. Rather, it is the hive that is the true animal, functioning according to its own biological necessity.
Just as a dog will shed its coat in summer, when a hive becomes overcrowded it will produce more queens. Nourished by the hive with royal jelly, the so-called swarm queen emerges from her queen cell and forces the incumbent queen whom the hive has denied food so she'll be light enough to fly to flee and start another hive with a contingent of field bees that follow her.
Once in possession of the hive and often after killing rival swarm queens the presumptive queen embarks on her maiden flight. High above the trees, the virgin queen mates with up to twenty drones. Their biological function complete, the drones fall away and die. The new queen then returns to the hive and begins laying eggs. Unless she is later usurped by another swarm queen, this is the only time she'll ever fly.
It is this predictable process that allows beekeepers like Bergman to "split" hives by taking half of the bees from a hive that is preparing to produce a swarm queen and placing them, via the hive box's removable honeycomb trays, into a new hive. The beekeeper then introduces a queen to the new hive, which in a few weeks will again grow to capacity effectively doubling the beekeeper's honeybee population.
"There's no other pollinator that we have such a handle on," says Berenbaum. "Bees have this social structure, and behaviors that go along with that social structure. With contemporary U.S. monoculture agriculture, we have vast expanses of crops that simultaneously need pollination services over a very short amount of time. There are not many insects that have the population capacity to meet that kind of demand. [In a single hive] there are 30,000 to 40,000 foragers at any given time."
Not only have honeybees been able to meet the population demand of an agricultural system based on vast tracts of the same crop blooming at the same time, they are also attractive because of their sheer versatility. Unlike, say, butterflies, which primarily pollinate wildflowers and only a few cash crops, honeybees can handle hundreds of different flower species.
The very versatility that makes the honeybee so attractive to modern agriculture has made it a highly managed species, susceptible to a host of genetic problems.
"[Before varroa mites] we had this background of feral bees. Queens would fly off and mate with whatever drones they found," explains the U of I's Charlie Whitfield. "In a colony you'd have a fair amount of diversity; all the workers would have the same mother, but they'd have genetically diverse fathers. With artificial insemination, they'll still have multiple fathers, but you're getting drones that are in your own apiary."
The lack of genetic variation isn't a problem, so long as a hive remains healthy. But when an unknown pathogen attacks, the hive may well lack the necessary genetic variants to adapt to the new threat. Often that means death for the entire hive.
What's more, while the social structure of the hive may be efficient at protecting young bees from outside toxins and pests, honeybees have roughly half as many detoxification genes as other insects, which might render them more susceptible to, say, manmade insecticides. But scientists don't know whether this is the case, nor whether the paucity of genes is a function of human intervention or natural evolution.
"They're not utterly incapable of processing pesticides otherwise we couldn't use them," Berenbaum notes. "But we have this confounding problem, in that people care about dead [honey]bees. They notice them. If there's an accidental insecticide drift that wipes out a population of carpenter bees, who's going to notice? Who's going to care? There's this problem of beekeepers pointing out every mass killing of honeybees so the perception is that they must be very susceptible to pesticides. But in reality we don't have an abundance of data."
Once he has extracted a bee's innards, Reed Johnson moves quickly through the crowded biology lab and places the digestive tract back into the deep freezer.
Like the rest of the CCD working group, Johnson is studying frozen bee samples that predate CCD, along with specimens culled from collapsed hives. One of CCD's primary characteristics is that adult bees leave the hive and do not return a knotty issue for researchers who are left with few options but to study the survivors.
"You can't do a lot of the genetic analyses on dead bees, because the part we're interested in degrades immediately," Johnson says. "You have to have live bees that were frozen on dry ice. So they're hard to get."
Once Johnson has collected enough gut samples, he places them into a test tube and pulverizes them with an electric pestle, then uses a centrifuge to extract the RNA from the mash. Like the rest of the CCD working group, Johnson is a beneficiary of geneticists who last year published the honeybee genome all 11,000 genes of it.
By using a microarray (a device that allows researchers to determine what genes were active at the time of death), Johnson will be able to discover the bee's biological state at the moment it was collected. If a sample of CCD bees were responding to, say, an ingested pesticide, those detoxifying genes would show up as active on the microarray.
"If the bee was in heavy-duty detoxification mode because it was exposed to some sort of a toxin, you'd expect to find many copies of genes involved in detoxification in that bee," Johnson elaborates. "If there are many, many copies of detoxification genes in the CCD bees, that would be evidence that it was possibly under some sort of environmental stress when it died."
But vast tracts of the insect's genetic code have yet to be cracked.
"That's the big problem," says Johnson. "Any conceivable genetic change should be picked up by the microarray, but the real problem is in figuring out what that means. Half of these genes we don't even know what they are."
Researchers at other labs, meanwhile, are looking into other potential culprits. Diana Cox-Foster, an entomologist at Pennsylvania State University who heads up the CCD working group, is investigating whether the plague may be the effect of a new or emerging pathogen. So far Cox-Foster's team is the furthest along of any group.
"We've identified a group of pathogens that are only associated with the CCD colonies and are not found in healthy colonies," Cox-Foster says. "We think that they are in large part responsible. But they're not completely responsible these pathogens are emerging themselves, and some of them are new to the United States."
Cox-Foster says her findings will be published in an upcoming issue of the journal Science. Citing the journal's customary press embargo on upcoming articles, she declined to discuss her findings in detail.
At any rate, even Cox-Foster's research remains preliminary. "Now we need to come back and demonstrate that indeed these are the causal organisms," explains the Penn State biologist. "We know that some of them have been found before. But they may have gone through some genetic changes, or been modified somehow, so we need to follow up on our research."
She is also quick to emphasize that pathogens are only partially responsible for CCD, which is almost certainly caused by a variety of modern stresses that afflict today's honeybees.
In the early days of the epidemic, it was suggested that cell phones might be affecting bees' navigational apparatus. Some beekeepers theorized that genetically modified corn was to blame. Various other culprits, including solar flares, al-Qaeda and even some sort of apian rapture, have been floated. One person blamed Kevin Federline, noting that bees had only gone missing in states where the artist had performed.
"The cell phones? We feel fairly comfortable that they're not playing a major role here," Cox-Foster says. "The corn we're fairly certain is not a major player." Ditto the sun, the terrorists and K-Fed.
Insecticides, though and specifically a class called neonicitinoids are a target.
"Although these are relatively safe for human consumption, they are very toxic chemicals to the pollinators," says Cox-Foster. "There's evidence that these chemicals can build up in the pollen and the nectar at sublethal concentrations for the bees, and that this may affect their overall physiology, including maybe their learning processes for instance, how well a bee can memorize its route when it leaves home."
But if insecticides are killing the bees, a larger question remains: Why now?
"There are clearly sublethal effects. Someone would have to find that there's been heavier use of those insecticides in the last year or so," says Jay Evans, an entomologist who works for the United States Department of Agriculture. "Insecticides don't spread the way a pathogen does. You have to really be in the area where it's been sprayed to show the effects."
Given their seemingly infinite replicability, honeybees will likely survive this latest assault, just as they have varroa and trachea mite, wax moth and hive beetle. Many believe the real danger is that beekeeping will cease to be an economically feasible profession.
Often passed down through families, it is a physically demanding trade that requies a strong knowledge base. Lose too many bees, and we may sacrifice that expertise.
"I'm really in a bad place here," says Glen Davis, a migratory beekeeper with a midsize operation in the western Missouri town of Bates City. Davis says that over the winter he lost 500 hives to CCD roughly 80 percent of his inventory. He spent nearly $25,000 on replacement bees, only to watch many of the new hives perish.
"I'm still trying to find out why the new bees that I put in there died," says Davis. "I've got to wonder if the rest of my bees are going to catch it, and if they're all going to die again."
Pulling into a tree-shaded clearing about a half-mile from the banks of the Missouri River, Sharon Gibbons steps out of her Ford F-150 pickup and locates a plank resting atop two poles that have been driven into the ground.
The Missouri River is flooding towns in western Missouri, and Gibbons, a 30-year veteran beekeeper who at age 66 has a head of curly gray hair and a motherly nature, has come to this small bee yard to move her hives to higher ground.
Stepping into a work-worn beekeeper suit, Gibbons places a yellow headband around her forehead to keep the sweat from her eyes as she moves her bees onto the elevated plank.
Along with her son, Chris, Sharon Gibbons produces honey, which she markets to area stores under the label Gibbons Bee Farm. The Gibbonses keep roughly 125 hives scattered throughout St. Louis County. They also have another, larger operation near Columbia.
The family's bees had a tough winter. Now, with the numbers down and the "honey flow" about to begin, there's no time to lose.
"Last summer was a bad drought and the honey production per hive was way down," Gibbons recounts as she slides a hive onto the makeshift platform. "So when we went into winter, the bees were already stressed out. A low honey-production year keeps the queen from laying the amount of eggs that she needs to have a strong enough hive to make it through the winter."
Before last winter Gibbons maintained about 1,000 hives. She estimates that she lost 20 percent to 30 percent of her St. Louis County stock and 60 percent of the bees in her Columbia operation.
"We have yards that were totally devastated," says Gibbons, who coos over her bees as she supplies them with sugar water, calling them her "girls." "It could have been CCD that dwindled the hive, but we don't know, because we've also got mites."
In other words, it's business as usual, and the work must go on. Today Sharon and Chris Gibbons are attempting to restore the population by splitting the stronger hives in half and adding new queens purchased from a Texas breeder.
"[We're] trying to get our numbers up, so that when everything blooms the bees will be able to capitalize on the honey flow," says Chris Gibbons, who tends to the Columbia hives.
Because the late-February frost that killed the area's elm and maple blossoms deprived the Gibbons' bees of the season's first shot of nectar, mother and son must put out sugar water literally, store-bought sugar mixed with water as they split their hives. With nectar (or in this case sugar water) plentiful, the queens will be encouraged to lay plenty of eggs, and the hive can double or even triple in size in a few weeks.
"We need to keep that queen laying. If she stops because there's no food, then you're setting back the hive," explains Chris Gibbons. "This doesn't provide them with much food, but it's enough to get them going and that's all we're looking for. Once they start bringing in nectar from the flowers, we'll stop feeding them."
Although Sharon Gibbons has dabbled in providing pollination services, her mainstay has always been honey. Now she says she's being approached by farmers in need of pollination services. Between last summer's drought and the winter's high mortality rate, she worries that she might not be able to get by as a beekeeper without becoming a pollinator.
"There are brokers who call me all the time. They'll take care of my bees from the time they leave my farm on the semi until they bring them back," she says. "But my experience with pollination is that the bees are not ready for the nectar flow when they come back."
As the day draws to a close, Gibbons moves on to north St. Louis County. It's coming on dusk, and as the beekeeper moves the hives, splitting them, cleaning them and rearranging them, her bees fill the air in an angry swarm.
"When the price of honey is way, way down, we can buy honey cheaper than we can produce," she says, gazing up through a cloud of bees. "Honey can be profitable but only if we don't keep on having all of these disasters."