compiled by Brian Tomasik
First published: 4 Jan. 2017; last update: 12 May 2017
This page collects information on the "cognitive" abilities of unicellular organisms.
Note: This page is a work in progress and doesn't have much diversity of sources yet.
See also: "Microbial intelligence"
Bray (2009), p. 18:
single cells are aware of their surroundings. They detect chemical flavors, mechanical vibrations, visual stimuli, electric fields, and gravity. They respond by moving or by changing their shape or internal state, selectively, in a discerning manner.
Bacteria exhibit the basic properties of living organisms, including growth and reproduction, environmental sensing, and responsive behavior.
Bryce Huebner notes:
I'm in the process of writing a tiny bit of a paper right now about bacterial behavior. It's in the context of a broader discussion of group behavior, and I realized that it was little more than bad ideology that leads philosophers to focus on complex prokaryotes in thinking about adaptive forms of group behavior. One of the things that keeps striking me as I read through the relevant literature is that the behavior of these tiny machines is really complex and highly adaptive. Their preferences might be simple, and they're probably grounded in little more than an impulse to pursue nutrients and avoid toxins, but bacteria seem to pass the William James test for mentality (fixed ends, with varying means). This is leading me to think that the whole idea of mindedness is poorly conceived, or that bacteria are minded...
This article quotes professor Jeffrey Lichtman:
Minds drum up the feeling of pain to keep the body safe, he continued, but the animal kingdom is full of self-preserving behaviors. Lichtman even suspected “that the first living animals that could affect their position in the world by moving very quickly developed ways to avoid putting themselves at risk, and that is equivalent to pain.” He and Dennett agreed that even microbes have incentives to sense their surroundings and avoid harm analogously to how humans avoid pain.
Bray (2009), pp. 5-7:
In 1854 the German biologist Wolfgang Pfeffer showed that if he introduced a capillary pipette filled with nutrient mixtures such as yeast or meat extract into a solution containing swimming bacteria, the bacteria would collect around the pipette and eventually enter into its tip. Capillaries filled with acid, alkali, or alcohol had the opposite effect, causing the bacteria to swim away. Other investigators at the time observed bacteria responding to light, temperature, or the concentrations of salts. [...]
Escherichia coli can detect something like fifty distinct chemicals. The list includes sugars and amino acids that act as attractants (the bacterium swims toward them) and a motley mixture of heavy metals, acids, and toxic substances that are repellents. E. coli's sensitivity is legendary. Even the slightest whiff of the attractant amino acid aspartate (a concentration of less than one part in ten million) is enough to change its swimming. [...]
to pursue any direction for more than a second or so, bacteria have to continually reassess their situation. How do they do this? The answer is that they have a sort of short-term memory that tells them whether conditions are better at this instant of time than a few seconds ago. By "better" I mean richer in food molecules, more suitable in acidity and salt concentration, closer to an optimum temperature, and so on. If on average conditions have improved, or at least are not any worse, then the bacteria will continue to swim in the same direction. But if conditions are deteriorating, then the bacteria tumble; they swim off in a new direction, selected more or less at random. The repeated execution of this pragmatic routine carries them over long distances and complicated terrains toward favorable locations.
This video says: "It's fair to say that E. coli has a short-term memory, but it's only about 4 seconds long. So what the cell does is it counts the things of interest in its environment over a second, looks up what it got for the previous 3 seconds, and then responds to the difference."
Freeman (2002) says (p. 274) regarding bacteria:
Although some species specialize by using just one type of food, the vast majority of bacteria are able to switch among several distinct sources of carbon and energy depending on which food items are available in the environment. [...]
[Intestinal E. coli] cells are capable of using a wide array of sugars to supply the carbon and energy they need. As your diet changes from day to day, the availability of different sugars in your intestine varies. Precise control of gene expression gives E. coli cells the ability to respond to these changes in its environment and use the different sugars.
Freeman (2002) also describes an experiment by Jacques Monod in the 1950s to test the hypothesis "that E. coli might actually prefer glucose to lactose as a food source" (p. 277). The experimental results were consistent with this hypothesis.
This video shows the behavior of many types of protists.
Applewhite and Morowitz (1967), pp. 332-33:
While protozoa lack the neurons of the metazoa, they do possess a fibrillar network that presumably may carry out the same functions (Kudo, 1966). Electron microscopy has been done on many of them (Pitelka, 1963), and they exhibit a wide variety of behaviors (Jennings, 1962, and Warden et at., 1940) that could be used to elicit learning. [...]
There have been many learning experiments with protozoa (McConnell, 1966, and Thorpe, 1963), but true learning in them has been questioned (McConnell, 1966).
Bichai et al. (2008), p. 510:
Free-living amoebae generally have 2 stages of development: the trophozoite and the cyst (Greub and Raoult 2004). The trophozoite is the active metabolic stage, feeding on bacteria and multiplying by binary fission. Hostile pH conditions, osmotic pressure, temperature, or even unfulfilled nutritional needs of the amoeba can cause its encystment (Greub and Raoult 2004).
In other words, amoebae can detect environmental conditions and respond adaptively to them.
Bray (2009), pp. 11-12:
A single cell of Amoeba proteus is a tiny speck of cytoplasm less than half a millimeter in length. Its amorphous, highly changeable form features fingerlike projections -- pseudopodia -- from a central mass. The cell lives on the bottoms of streams and ponds and typically crawls over the surfaces of pebbles and vegetation by extending and retracting its pseudopodia. When it encounters obstacles in its path, regions in contact with the obstacle become quiescent while those farther away move more actively. Consequently the cell flows in a new direction. Other modes of movement are also seen. For example, a hungry amoeba often adopts a more rapid form of progression termed loping, alternately extending and contracting its body like an inchworm. Another state I like to think of as skydiving occurs when an amoeba is freely suspended in water. The cell now adopts a starlike form with long pseudopodia extending in all directions. It seems to be searching for a suitable surface. Indeed, no sooner does one pseudopodium make an effective contact than currents of cytoplasm flow in that direction. Other pseudopodia retract; the cell settles onto the surface and resumes its normal mode of creeping.
[...] Herbert Jennings, a naturalist working at the Zoological Laboratory of the University of Michigan in the late nineteenth century, observed spherical cysts, or spores, of Euglena (a ciliated protozoan) being used as food [by Amoeba proteus]. The amoeba in his cultures attempted to engulf the cysts by extending lateral pseudopodia and a thin sheet of cytoplasm. If the attempt was successful, then the cyst was enclosed in a cup-shaped depression, entered the cell in a parody of swallowing, and was eventually digested. The cysts, however, were smooth and rolled easily when touched. Jennings recorded an amoeba that pursued a rolling cyst in a circular path for fifteen minutes, making repeated attempts at capture, until the cyst was eventually lost. "This behavior," Jennings remarked "... makes a striking impression on the observer who sees it for the first time. The Amoeba conducts itself in its efforts to obtain food in much the same way as animals far higher in the scale."
Rather surprisingly, Amoeba proteus is also able to capture actively swimming organisms such as the ciliated Paramecium despite its lumbering gait. It seems to be able to detect swimming paramecia at some distance and may pursue them over long distances. Once contact is made, the amoeba initiates a slow encircling action and attempts to trap the struggling prey. If encounters with swimming prey are sufficiently frequent, the amoeba anticipates a capture by forming a cup-shaped depression on its surface like a baseball mitt.
Bray (2009), p. 18:
An amoeba can immediately tell a Euglena cyst from a grain of sand of the same size and will devour the former while rejecting the latter.
While Stuart Hameroff's views on consciousness are dismissed by most scientists, Hameroff does make an important observation about the abilities of unicellular organisms that lack nervous systems:
A single celled paramecium swims around. It finds food. It finds a mate. It has sex. It can learn. It can avoid predators. It can avoid obstacles. It's one cell. It does it with microtubules.”
Wikipedia says that Stentor "are among the biggest known extant unicellular organisms."
Video: "Stentor hunting for food".
Bray (2009), pp. 14-17:
Unexpectedly, the most intricate behaviors so far recorded in unicellular animals have been found in organisms attached to a surface rather than swimming free. The trumpet-shaped Stentor spends most of its time attached to water plants or debris by an irregularly shaped foot embedded in a transparent tube of mucus. Its slender body and flattened, discoid mouth cavity are lined with fine cilia. Beating of these cilia sets up continual currents of fluid that carry particles of suitable size into the mouth cavity. The animal then decides whether they are edible or not. Particles of food are retained and ingested, whereas indigestible particles are ejected. The organism can also turn or sway on the end of its long stalk, thereby pointing its mouth cavity in selected directions.
The lynx-eyed Herbert Jennings gave a detailed account of the response of Stentor roeselii to experimental stimulation. A fine current of water squirted at the open disk caused the cell to contract in a flash into its tube. After about half a minute the animal extended again and the cilia resumed their activity. A second application of the jet of water identical to the first, however, was then ignored, and the animal continued with its feeding activities. Jennings observed a similar sequence of a "surprise" response followed by rapid acclimatization with a whole variety of stimuli, such as a small jarring of the animal's container. Once again the organism displayed a simple form of memory.
Jennings found, however, that a potentially injurious stimulus, such as a cloud of particles of the red dye carmine introduced to the feeding disk, evoked a richer set of actions. The stentor at first makes no response, taking the obnoxious particles of carmine into the pouch and into the mouth. After a short time, though, it turns to one side by bending its long stalk (always in the same direction) as if to move its mouth out of the path of the cloud of noxious particles. This may be repeated: the animal twists on its stalk two or three times about its long axis. If the repeated turning to one side does not relieve the irritation, a second strategy is tried. Beating of the ciliary hairs on the body is suddenly reversed in direction, thereby causing particles in the disk and in the pouch to be ejected. The reversal lasts but an instant before the usual current is resumed. If the irritant particles continue to arrive, the reversal is repeated two or three times in rapid succession.
If stentor still does not get rid of the stimulation in either of the ways just described, it contracts into its tube. In this way, of course, it escapes the offending cloud, but at the expense of losing all opportunity to obtain food. The animal will now remain in the tube about half a minute before again cautiously extending. When the body has reached about two-thirds its original length, the cilia around the disk-shaped mouth begin to unfold. They recommence their beating, allowing the animal to recommence feeding.
What happens now? Suppose that the water currents again carry noxious carmine grains into the feeding disk. The stimulus and all the external conditions are the same, so will the stentor go through the same routine -- bend to one side, reverse its cilia? No. The animal has become changed by its experience and tries a new strategy. This time, no sooner do the carmine particles reach its disk than the animal contracts fully into its tube. The sequence may be repeated many times, the animal cautiously extending after a while and, if it still finds particles, contracting again. Each time, it stays a little longer in the tube and seems to be more reluctant to emerge. Finally, it contracts violently and repeatedly while still enclosed in its tube, so violently that the foot attachment breaks and the animal is set free. Stentor leaves its tube and swims away, avoiding if necessary the cloud of carmine and other obstacles, and sets sail for a new home.
In due course, the minute creature will settle onto the bottom of its pond or container. It then behaves in a peculiar way. The partially unfolded disk (the part previously used for feeding) is now used as a means of locomotion. Stentor now creeps rapidly over the terrain, following all the irregularities of the surface rapidly and seeming to explore it thoroughly. This may last for sometime, the animal often quitting one site and swimming on to another, exploring heaps of debris and surfaces of solids in its path.
After ~10-20 minutes, the stentor decides on a location, builds a new mucus tube for itself to live in, and attaches to its new home (Bray 2009, p. 17).
Bray (2009), p. 18:
Most microorganisms display what in higher animals is termed attention. A stentor or amoeba dislodged from a surface actively seeks a new site of attachment. While engaged in this search it ignores other stimuli such as changes in temperature or chemical signals that produce an immediate reaction in a free-living individual.
Bray (2009), p. 18:
Almost all single-celled organisms respond to light. Ciliates such as paramecia move to weak light but are repelled by strong light. The single-celled alga Chlamydomonas has a so-called eyespot containing pigment molecules that enable it to detect the direction of incoming light.
Computation by single neurons
A single neuron of a multicellular animal is a fairly complex computing device in its own right. One example of this point can be seen from the field of "dendritic computing".
Senior author Professor Michael Hausser commented: “This work shows that dendrites, long thought to simply ‘funnel’ incoming signals towards the soma, instead play a key role in sorting and interpreting the enormous barrage of inputs received by the neuron. Dendrites thus act as miniature computing devices for detecting and amplifying specific types of input.["]
London and Häusser (2005) review the topic of dendritic computation. Here's one tidbit from the paper (p. 512):
Each subunit thus integrates its inputs and passes them through a [sigmoidal] nonlinearity function. This gives each piece of dendrite the computational power of a small unit similar to those conventionally used in neural networks. The output of each subunit is conveyed to the soma, in terms of passive dendritic integration. The picture that emerges from this analysis is of a two-layered neuronal network that resides within a single neuron. This analysis is supported by detailed modeling of single neurons, showing that the predictive power of the two-layer neural network description is very good (Poirazi et al. 2003b), and by experiments in layer 5 pyramidal neurons (Polsky et al. 2004). The attractiveness of this approach stems from the fact that two-layer neural networks are general-purpose computation machines, which have been extensively studied and can implement very powerful computations.