Monday, June 10, 2013

House-hunting the democratic way - learning from the bees...





















When I walked into the lecture hall last week, this image looked like a swarm of bees to me. By the time, I walked out one hour later, this is what I saw.


No, this was not the effect of a dream state or a hallucinogen but rather the effect of Dr. Thomas Seeley’s work.

Tim Seeley, as he is known, is an ethologist and an entomologist from Cornell and probably one of the world’s leading expert in the phenomena of swarm intelligence. He has been studying the behaviors and social structures of bees for many decades now. As far as scientific genealogy goes, Tim has great Pedigree. He began his work on bees, as part of his doctoral dissertation with Lindauer who was a student with Karl Von Frisch, the Nobel laureate who pioneered the field itself. It was Von Frisch’s discovery of the now famous waggle dance in bees, as a tool for communication that gave us our first insights into the social structures of the insect world. Von Frisch found that scout bees would return from a food source and would then dance around the hive; soon after which, the rest of the forager bees would go hunting and collecting.


By careful observation over years, Von Frisch found that the duration, direction and quality of the dance would communicate to the rest of the hive, details about the quality, location and distance of the food source. This opened up a whole new field in biology as animals were long considered to be incapable of communication, coordination, culture and altruism. Instead, Von Frisch’s findings attacked the very foundations of human uniqueness and demonstrated these tiny insects to be capable of all this and much more.

Lindauer, Von Frisch’s student and later colleague, followed up on this work and found that a young bee swarm would also choose its future nest site by a decision making process that was based on reports of the dancing scouts. Seeley was intrigued by the idea of swarm intelligence, by the ability of a swarm as a whole to arrive at a decision that could potentially make or break the future of the colony.

His first forays into this question began with determining what the bees were looking for, when they were scouting the real estate market. Honeybee colonies reproduce by budding, whereby the queen and some workers (a few thousand) leave the nest and bivouac on a branch for a few days depending on their resources. At this point, the swarm needs to locate a new home for themselves that will ensure their survival. By chasing bee swarms and by building potential hive sites of many different shapes and sizes as part of his doctoral work, Tim found that the young swarm was looking for a nesting site that was high, well insulated and yet roomy. It had to be high enough to escape predators (at least 15 ft), big enough (atleast 10 gallons) to rear a new brood and to stock supplies to last through the winter. For a honeybee swarm, choosing a potential home is a momentous decision that might compromise their survival itself. The task of hunting is thus delegated to the most experienced workers who scout nearby locations and report to the swarm. 

Lindauer had already shown that the scouts used the waggle dance to “tell” the rest of the swarm about the new site and that the decision was somehow arrived at by a collective process. Trying to understand the dynamics of the swarm was a challenging and almost impossible task at the time, even for someone like Seeley. In the early decades of the field, ecologists were limited to the use of a simple tool set of notebooks, pens, stopwatches, paint sets, wristwatches and little else. Tracking hundreds of individual scouts, their movements, dances and patterns at the level of multiple swarms was impossibility. Even someone as creative and ingenious as Seeley was forced to wait for technology to catch up with his ideas.

Finally, in the last decade of the twentieth century, as the camera and video recording techniques came of age, Seeley decided that the time was ripe for delving into those forbidden questions. And the answer lay in careful and meticulous observation. Seeley and his students would manually label and mark the thousands of bees in a swarm, but this was only the easy part of the job. They would then video record the entire house-hunting process for a period of 2-3 days and at the end of it, they faced the enormous task of decoding the videos to identify the underlying patterns. The watched the scouts hunt at different locations, coming back to report and then the entire decision making process. Each 16 hour film of the bees’ decision making process would take atleast a month of eye-stinging labour to decipher. What they found was fascinating indeed.

The experienced scout bees would head out and survey the available real estate. They would the, come back and advertise the potential locations and their qualities to their nest mates by performing the waggle dance. The angle of the bees with respect to the sun would indicate the direction of the site, while the duration of the dance would indicate the distance. The quality of the site was indicated by the simple metric of the number of dance repeats that the bee did. The more a scout bee advertised, its location, the stronger the lobby it built. These waggle dances would then recruit additional scouts to the site until a decision was made. Interestingly, each scout bee would only visit one site on most occasions and would only lobby or rather “dance” for it. Each time she returned from the site, the number of dance circuits would progressively decline but each routine would recruit a new wave of scouts, creating multiple, independent reports. 


Seeley and his students began their studies suspecting a consensus building mechanism to be involved – a democracy of sorts. What they instead found was that the scouts themselves don’t pay any attention to the consensus. They decision is made by a quorum, a critical number (20-30) of bees being simultaneously present in the nest site – representing successful reports from many independent scouts. Trying to understand the significance of the number, Seeley and Dr. Kevin Passino, a professor of computer engineering at the Ohio state university, came together and modeled this decision making process. They fund that in a simulation, adjusting the quorum to only 15 bees would result in quick but error prone decisions. Increasing the number to above 20 on the other hand produced slower but only slightly more accurate decisions. It thus seemed that the bees had found the optimum number to balance the cost of resource starvation with expected gain in accuracy. But the precise mechanism underlying this “plebiscite” remained unclear, until of course, Dr. Seeley and his students heard the head butts.

They found that in deciding which site to choose, the scouts would also employ another tactic in addition to the positive-reinforcements achieved by dancing. They also had a stop signal – a booming head butt. When choosing between two nest sites, the scout bees committed to each nest site would direct these head butts to the scouts promoting the other box; thus setting up a cross inhibition between the two populations of scout bees.

The honeybee stop signal is a vibrational signal signal that lasts about 150 ms and has a fundamental frequency around 350 Hz. It is typically delivered, by the sender butting her head against the dancer. Although the dancer may not show an immediate response to the signal, the accumulation of such stop signals is seen to increase the probability that the bee will cease dancing. [Commonly used during foraging, the stop signal is given when a forager bee is attacked in a food source and this reduces the recruitment of colony resources to perilous food sources.]


To further understand the implications of such a cross-inhibitory selection mechanism, Dr. Seeley and his colleagues further modeled the decision making process. Testing individual models of no or indiscriminate stop signals suggested the overwhelming possibility of reaching at an impasse – an impossible deadlock. Such a stable deadlock would result in the swarm never reaching a decision and thus starving to death. Simulations showed that the deadlock persists even when the stop signaling is applied with discretion but below a critical threshold.

When the stop signals are applied above a critical threshold, the swarm is either able to randomly arrive at a consensus for one the two equal alternatives or if the difference in the quality of the two sites is significantly different, then the stop signals ensured a consensus for the superior alternative. The existence of the stop signal and a high enough quorum thus ensure that the bees arrive at an optimal decision most of the times.

Once the search committee has made up its mind and the critical quorum of bees is established at the nest site, the scouts lobbying for the site begin their final “move”. They begin with an auditory signal, called as worker piping, that can actually be heard by us. Coincident with this signal, the other bees in the swarm begin to prepare for flight. Bees fly with wings moving at some 250 times per second and this requires their wing muscles to be warm (35 deg C) and metabolically active. To do this, upon receiving the piping signal, the bees warm up their flight muscles by disengaging their chest muscles from their wings and vibrating them. The temperature of the whole swarm rises up rather steeply in less than half an hour.
 
Worker piping and the pitch of the sound perfectly match the beat frequency of a flying bee and some early experiments suggest a critical link between the two. In the final few minutes before the take off, the workers exhibit another rather strange phenomenon - a so called ritualistic buzz run. Here, the bees run across the entire swarm with outspread wings and a buzzing sound, almost invoking everyone into action.  As the swarm is warm and ready, the buzz run begins and finally the swarm lifts-off. It slowly hovers over its temporary home for a minute or two and then starts proceeding towards the nest at speeds ranging from 0.5 – 5 miles per hour. They finally stop near the goalm their future home scented with the Nasonov pheromones from the scouts. In the absence of these scent markers, the swarm finds its more difficult, not impossible, to locate the entrance to the hive site. At this point, they are then guided by these streaker bees, who repeatedly shoot towards the goal and guides the others. The bees thus finally make it to their new home – and a young hive begins to grow.

Although elegant and simple, this mechanism needs further careful study to account for the individuality and subjectivity of the bees.  If in the early phase of the scouting, a single bee dances more enthusiastically for a bad site or less enthusiastically for a good site, then the balance of signals could tilt very rapidly introducing a lot od stochasticity. In fact, as they individually tracked these bees, Dr. Seeley and his group also noticed many differences between them. They are all not the same like a cookie cutter and are rather quirky. Some are really good dancers, some are not; some are really peppy and get going early in the morning while some others have to be woken up. They have a lot of personality and with the cumulative mechanism of this process, small vulnerabilities at the start could easily run amock and alter the fate of the hive.

It would also be interesting to know if the hive changes its criteria and its choices if they are running out of reserves and how the choices are made? The rate of information arriving to the hive could also bias the processing of the input and the cascade to the decision – say a scout flies closer to the swarm and reports back sooner and builds up a bigger lobby. The reliability of the responses of the individual bees to multiple optimal or suboptimal sites is another interesting question that may be rather revealing. All in all, like all great studies in science, Dr. Seeley’s raises many more interesting questions, even as he finds the answers to them.

Interestingly, as Dr. Seeley and his students were uncovering these details of the house-hunting process of the swarm, there emerged a few parallels that are imperceptible to an otherwise untrained mind. What Seeley saw was an emerging parallel between the bee swarm and the human brain. In a swarm, each scout reported on a single find, much like a neuron responding to a particular stimulus. “Both are cognitive entities shaped by natural selection to be skilled at acquiring and processing information to make decisions,” he says. In both these systems, decision-making is a competition between mutually interacting populations of excitable units – neurons or individuals – that accumulate noisy evidence for alternatives and, when population exceeds a threshold, the corresponding choice is made. A common underlying feature for both bees and neural decision-making is the existence of cross-inhibition. A population that inhibits others proportional to its own activation, thus ensuring that only one of the alternatives is chosen.

And so, as I stepped out of that lecture hall at the end of that hour, my rewired brain could see more to the statement “buzz in the brain” than was ever obvious to me. 

The question is what do you see?




References:

1)   Stop signals provide cross inhibition in collective decision making by honeybee swarms, Seeley et al, Science, January 2012
2)   Swarm intelligence in honeybees: Talk by Dr. Tim Seeley at the University of California, San Diego.
3)   Decoding the language of the bee: Nobel lecture, Dec 12, 1973, Karl Von Frisch.
4)   Swarm Intelligence: How Tom Seeley discovered ways that bee colonies make decisions, Part II, MEA McNeil
5)   How honeybees break a decision-making deadlock, Science, 6 January 2012, By Jeremy E Niven
6)   You tube Lecture by Dr. Tim Seeley

 









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