The characteristic extinction pattern which is observed when the submucosa is viewed in the optical polarizing microscope has been analyzed in terms of the configuration and orientation of the 4 micron diameter collagen fibers. It is shown that the observed polarization effects are produced by periodic variations in orientation of fully birefringent fibers. The fiber configuration required to produce the observed polarization effects is a tilted wave configuration with a crimp period of approximately 20 micron. In the model, the tilted waveform fibers are crimped in register and form parallel arrays. The arrays are oriented in layers at approximately +30 degrees and -30 degrees to the longitudinal direction and are mirror images of each other. Analysis of the extinction pattern shows that the model satisfactorily accounts for the observed polarization effects at several different angles of the crossed polaroids. The calculated strain necessary to straighten the wavy fibers of the model correlates well with the observed strain to uncrimp the collagen fibers in the intestine. This suggests that the initial response to stress is gradual uncrimping of the collagen fibers, and concurrently, a decrease in the angle between biaxially oriented fibers, rather than extension of the straight fibers.
Collagen fibers of rat intestine were observed with and without mechanical stress in the scanning electron microscope. Observations were correlated with the previous results obtained by optical polarized microscopy to provide further insight into the organization and morphology of intestinal collagen. Larger fibers, approximately 4 micrometers in diameter, are densely packed in parallel undulating arrays. The initial response to stress is straightening of the original fibers. The extended fibers are biaxially oriented at +30 degrees and -30 degrees to the longitudinal direction. These large fibers appear as assemblies of subfibers. At higher magnifications, the larger fibers appear to be enmeshed in a network of small randomly oriented fibers approximately 0.2 micrometer in diameter. Study of the effect of age on fiber morphology showed that the length of the large fiber undulations increases during maturation but remains constant during aging. The diameters of large and small fibers appear not to change with age, but more of the 4 micrometers fibers are loosely associated into larger fibers which can be observed at both the optical microscope and scanning electron microscope levels. A hierarchical organization of intestinal collagen is proposed.
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