At the ultrastructural level, epithelia performing solute-linked water transport possess long, narrow channels open at one end and closed at the other, which may constitute the fluid transport route (e.g., lateral intercellular spaces, basal infoldings, intracellular canaliculi, and brush-border microvilli). Active solute transport into such folded structures would establish standing osmotic gradients, causing a progressive approach to osmotic equilibrium along the channel's length. The behavior of a simple standing-gradient flow system has therefore been analyzed mathematically because of its potential physiological significance. The osmolarity of the fluid emerging from the channel's open end depends upon five parameters: channel length, radius, and water permeability, and solute transport rate and diffusion coefficient. For ranges of values of these parameters encountered experimentally in epithelia, the emergent osmolarity is found by calculation to range from isotonic to a few times isotonic; i.e., the range encountered in epithelial absorbates and secretions. The transported fluid becomes more isotonic as channel radius or solute diffusion coefficient is decreased, or as channel length or water permeability is increased. Given appropriate parameters, a standing-gradient system can yield hypertonic fluids whose osmolarities are virtually independent of transport ratp/'over a wide range, as in distal tubule and avian salt gland. The results suggest ~ that water-to-solute coupling in epithelia is due to the ultrastructural geometry of the transport route.T h e purpose of this paper is to examine whether some distinctive physiological properties of epithelia might arise as geometrical consequences of epithelial ultrastructure.Most epithelia absorb or secrete specific fluids, such as bile, the glomerular filtrate, gastric juice, aqueous h u m o u r , sweat, and cerebrospinal fluid. T h e p r i m a r y transported fluid m a y be either isotonic or hypertonic to plasma (hypotonic secretions in vertebrates appear to result from isotonic secretion ~o6i
In the modern world, biotic diversity is typically higher in low-latitude tropical regions where there is abundant insolation (light and heat) and low thermal seasonality. Because these factors broadly covary with latitude, separating their possible effects on species diversity is difficult. The Eocene was a much more equable world, however, with low temperature seasonality extending into lower-insolation higher, cooler latitudes, allowing us to test these factors by comparing insect species diversity in (1) modern, temperate, low-insolation, highly seasonal Harvard Forest, Massachusetts, U.S.A., 42°29'N; (2) modern, tropical, high-insolation, low-seasonality La Selva, Costa Rica, 10°26'N, and; (3) Eocene, temperate, low-insolation, yet low-seasonality McAbee, British Columbia, Canada, above 50°N paleolatitude. We found insect diversity at McAbee to be more similar to La Selva than to Harvard Forest, with high species richness of most groups and decreased diversity of ichneumon wasps, indicating that seasonality is key to the latitudinal diversity gradient. Further, midlatitude Eocene woody dicot diversities at McAbee, Republic (Washington, U.S.A.), and Laguna del Hunco (Argentina) are also high, similar to modern tropical samples, higher than at the modern midlatitude Harvard Forest. Modern correlations between latitude, species diversity, and seasonal climates were established some time after the Eocene.
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