The intrinsic complexity, variety of concepts and numerous ways to quantify landscape heterogeneity (LH) may hamper a better understanding of how its components relate to ecological phenomena. Our study is the first to synthesize understanding of this concept and to provide the state of the art on the subject based on a comprehensive systematic literature review of 661 articles published between 1982 and 2019. Definitions, terminologies and measurements of LH were diverse and conflicting. Most articles (534 out of 661) did not provide any definition for LH, and we found great variation among the studies that did. According to our review, only 10 studies measured the effects of different land-cover types on biotic or abiotic processes (functional LH). The remaining 651 studies measured physical attributes of the landscape without mentioning that different land-cover types may impact biotic and abiotic processes differently (structural LH). The metrics most frequently used to represent LH were the Shannon diversity index and proportion of land-cover type. Most metrics used as proxies of LH also coincided with those used to represent non-heterogeneity metrics, such as fragmentation and connectivity. We identify knowledge gaps, indicate future perspectives and propose guidelines that should be addressed when researching LH.

Membranous (M) cells in follicle-associated epithelium (FAE) play an important role in the mucosal immunity through transport of a variety of foreign antigens to the underlying mucosa-associated lymphoid tissue (MALT). We aimed to investigate the ultrastructure of M cells in the FAE covering nasal-associated lymphoid tissue (NALT) both in specific pathogen-free (SPF) rats and in conventional environment-adapted (SPF-CV) rats aged 8–38 wk. In NALT of both SPF and SPF-CV rats, FAE included the nonciliated microvillous cell, which appears to be an analogue of M cell previously described in other MALT. In SPF rats, M cells increased in number only slightly with age, and they maintained morphological uniformity irrespective of age. In SPF-CV rats, M cells selectively increased in number resulting in prominent expansion of FAE surface area in parallel with the duration of maintenance in a conventional environment. In addition, M cells in SPF-CV rats showed heterogeneity in their surface morphology such as the length and number of microvilli and cell surface area and outline. In addition, the FAE was stratified by various subtypes of M cells, which were characterised by several subcellular alterations including the presence of many keratin filaments, homogeneous dark bodies and extensive cytoplasmic interfoliation with wide intercellular spaces filled with amorphous proteinaceous material. These characteristics of M cells in SPF-CV rat were intimately related with a preferential influx of immunocompetent cells into the FAE, which was not seen or was very rare in SPF rats irrespective of age. The results suggest the possibility that NALT may effectively carry out the mucosal immune response against antigenic stimuli of different magnitude through the unique dynamics of M cells which seem to be influenced by the infiltration of immunocompetent cells.