The meaning of "degeneration" was poorly defined, but can be described as an organism's change from a more complex to a simpler, less differentiated form, and is associated with 19th-century conceptions of biological devolution. In scientific usage, the term was reserved for changes occurring at a histological level – i.e. in body tissues. Although rejected by Charles Darwin, the theory's application to the social sciences was supported by some evolutionary biologists, most notably Ernst Haeckel and Ray Lankester. As the 19th century wore on, the increasing emphasis on degeneration reflected an anxious pessimism about the of European civilization and its possible decline and collapse.