The metabolic rate and its scaling relationship to colony size were studied in the colonial ascidian Botrylloides simodensis. The colonial metabolic rate, measured by the oxygen consumption rate (V O 2 in millilitres of O 2 per hour) and the colony mass (wet weight M w in grams) showed the allometric relationship (V O 2 = 0.0412 M 0 .79 9 w . The power coefficient was statistically not different from 0.75, the value for unitary organisms. The size of the zooids and the tunic volume fraction in a colony were kept constant irrespective of the colonial size. These results, together with the two-dimensional colonial shape, excluded shape factors and colonial composition as possible causes of allometry. Botryllid ascidians show a takeover state in which all the zooids of the parent generation in a colony degenerate and zooids of a new generation develop in unison. The media for connection between zooids such as a common drainage system and connecting vessels to the common vascular system experienced reconstruction. The metabolic rate during the takeover state was halved and was directly proportional to the colonial mass. The scaling thus changed from being allometric to isometric. The alteration in the scaling that was associated with the loss of the connection between the zooids strongly support the hypothesis that the allometry was derived from mutual interaction among the zooids. The applicability of this hypothesis to unitary organisms is discussed.

The allometric scaling of metabolic rate of organisms, the three-quarters power rule, has led to a questioning of the basis for the relation. We attacked this problem experimentally for the first time by employing the modular organism, the ascidian that forms a single layered flat colony, as a model system. The metabolic rate and colony size followed the three-quarters power relation, which held even after the colony size was experimentally manipulated. Our results established that the three-quarters power relation is a real continuous function, not an imaginary statistical regression. The fact that all the hypotheses failed to explain why the two-dimensional organism adhered to the three-quarters power relation led us to propose a new hypothesis, in which the allometric relation derives from the self-organized criticality based on local interaction between modulus-comprising organisms.

SUMMARY In order to characterize the energy expenditure of Paramecium, we simultaneously measured the oxygen consumption rate, using an optic fluorescence oxygen sensor, and the swimming speed, which was evaluated by the optical slice method. The standard metabolic rate (SMR, the rate of energy consumption exclusively for physiological activities other than locomotion)was estimated to be 1.18×10–6 J h–1cell–1 by extrapolating the oxygen consumption rate into one at zero swimming speed. It was about 30% of the total energy consumed by the cell swimming at a mean speed of 1 mm s–1, indicating that a large amount of the metabolic energy (about 70% of the total) is consumed for propulsive activity only. The mechanical power liberated to the environment by swimming Paramecium was calculated on the basis of Stokes' law. This power, termed Stokes power, was 2.2×10–9 J h–1 cell–1, indicating extremely low efficiency (0.078%) in the conversion of metabolic power to propulsion. Analysis of the cost of transport (COT, the energy expenditure for translocation per units of mass and distance) revealed that the efficiency of energy expenditure in swimming increases with speed rather than having an optimum value within a wide range of forced swimming, as is generally found in fish swimming. These characteristics of energy expenditure would be unique to microorganisms, including Paramecium, living in a viscous environment where large dissipation of the kinetic energy is inevitable due to the interaction with the surrounding water.

During the spawning process in starfish, oocytes are arrested at metaphase of meiosis I (MI) within the ovary, and reinitiate meiosis only after they have been released into the seawater. However, this arrest does not occur if the ovary is removed from the animal. As the pH of the coelomic fluid is buffered by CO2/H(+)/HCO3(-), we investigated the involvement of gas concentrations in MI arrest. In vivo, the CO2 level in the coelomic fluid was high (∼1.5% vs. 0.04% in air) and the O2 level was low (0.1-1.0% vs. ∼20% in air). When these gas conditions were reproduced in isolated coelomic fluid or seawater, ovarian oocytes arrested at MI, just as in vivo. Isolated oocytes from the ovary required the similar high CO2 and low O2 level to remain arrested in MI and had an intracellular pH of ∼6.9. Intracellular pH increased to ∼7.3 when oocytes were transferred to seawater equilibrated with air, a condition that mimics that of spawning. We used ammonium acetate to clamp intracellular pH at different levels and found that MI arrest occurred when intracellular pH was ∼6.9. Our results support the idea that high CO2 and low O2 in the ovarian environment lead to low intracellular pH and MI arrest, while spawning into the seawater with low CO2 and high O2 results in high intracellular pH and release from MI arrest. The biological significance of MI arrest is that oocytes are spawned into seawater at the optimal physiological state of MI when the least polyspermy occurs.