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ULTIMATE COMPUTING - Quantum Consciousness Studies

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Models of Cytoskeletal Computing 159<br />

(1965) also proposed that receptor cilia operated “by virtue of transmitting<br />

mechanical signals to the base of the cilium.”<br />

Biologist Jelle Atema (1973) of the Wood’s Hole Oceanographic Institute,<br />

whose work had focused on acoustical perception at the cellular level, also linked<br />

microtubules and sensory transduction. Noting common cilia-like structure in<br />

sensory receptor organelles among a wide variety of organisms, Atema proposed<br />

that sensory cilia conveyed environmental information to the rest of the cell. He<br />

suggested that transduction occurred by propagated conformational changes in the<br />

microtubule subunits which constituted these cilia. He argued that microtubules<br />

were active functional units in reception and transduction of sensory information.<br />

Atema reviewed conformational changes in MT tubulin subunit dimers<br />

observed by a variety of authors and suggested that these occurred under<br />

physiological conditions. For example, oscillations of sperm flagella are<br />

apparently not controlled by their cell membrane but rather are direct properties of<br />

their microtubules in the presence of ATP (Lindemann and Rikmensboel, 1972).<br />

Atema concluded that a sequence of subunit conformational changes is likely to<br />

occur in microtubules. In motor cilia and flagella, the distortion would originate at<br />

the base of the structure and be propagated distally, resulting in wavelike or<br />

whiplike motions which propel the organism. In sensory cilia, the distortion<br />

apparently originates near the distal end of the ciliary MT and propagates in either<br />

direction or at least proximally towards the cell body. There the signal may<br />

propagate via either an excitable membrane or through the anchoring basal body<br />

and cytoskeleton. Because mechanical deformation and/or local chemical<br />

distortion are sufficient stimuli to start a propagating signal in sensory cilia,<br />

Atema’s microtubule theory assumed that distortion of tubulin conformation by<br />

any number of sources was sufficient to propagate a conformational wave. Thus<br />

light energy, chemical bond energy, and mechanical forces could be transduced<br />

by sensory cilia.<br />

Atema’s view of MT information processing was an “all or none”<br />

propagation by allosteric conformational changes along tubulin protofilaments.<br />

His was the first theory to look beyond the global behavior of cilia to consider<br />

conformational effects in tubulin components. Subsequent theories became more<br />

elaborate to consider localized analog functions, switching, and collective<br />

neighbor interactions among MT subunits.<br />

8.2.2 MT Mechano-lonic Transducers/Moran and Varela<br />

Biological sensory systems ranging from human inner ears to honey bee<br />

hairplate receptors use sensory cilia to transduce mechanical deformation into<br />

cognitive information like position in space (“proprioception”). Harvard<br />

biologists Moran and Varela (1971) studied the proprioceptive transduction of<br />

mechanical deformation in the legs of cockroaches. There, tactile spines contain<br />

mechanoreceptor structures called “campaniform sensilla” which consist of a<br />

single bipolar neuron from whose dendritic tip extends a modified cilium<br />

containing 350 to 1000 microtubules. Moran and Varela determined that this<br />

parallel bundle of MT was directly involved in the transduction of mechanical<br />

deformation to the neuron.<br />

All neurons are mechanical receptors, being stretch sensitive to cause ionic<br />

currents. In the cockroach campaniform sensilla, the cilium is the only bridge and<br />

transduction does not occur when MT are incapacitated with tubulin binding<br />

drugs like colchicine and vinblastine (Moran and Varela, 1971). MT are thus<br />

directly involved in proprioceptive determination of the system’s spatial relation<br />

to its environment. Moran and Varela saw two possible roles for MT in their<br />

mechanoreceptor function: as passive translation rods, or as generators of

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