Anatomical characteristics of roots of citrus rootstocks that vary in specific root length.

Among citrus rootstocks, higher specific root length (SRL, root length/d. wt) has been linked to several specific morphological and physiological traits, including smaller average root diameter, higher root hydraulic conductivity and higher rates of root proliferation. In this study, thickness of the outer tangential exodermal (hypodermal) wall and its suberin layer, number of passage cells, presence of epidermis, and stelar anatomy were examined and related to variation in root diameter of field roots of known maximum age. We also compared root morphology and anatomy of young roots in the field with those of potted rootstock seedlings in the glasshouse. Fibrous roots were measured separately from pioneer (framework) roots. Among the fibrous roots, only the first-order (root links having a root tip) and second-order (root links bearing first-order roots) laterals were examined. Among first-order field roots, larger root diameter was caused by larger rather than more numerous cells in the cortex. Root diameter of first-order roots was positively correlated with both number of passage cells in the exodermis and thickness of the secondary walls of the exodermis in both field and potted plants. Exodermal walls were about 80% thicker in field- than pot-grown roots. In the field, in more than 50% of the first-order roots examined less than 30% of the root surface was still covered by epidermis, with few differences among rootstocks. In contrast, in roots of 19-wk-old glasshouse plants generally 70-100% of the epidermis was still intact. There was no evidence of secondary xylem development in second-order fibrous roots in the field; in seedling, pot-grown rootsystems, 75-97% of second-order roots had formed secondary xylem despite their small diameter (<0.8 mm). It is argued that there can be suites of physiological, morphological and anatomical traits in roots that co-vary with specific root length. Investigations of how root morphology and anatomy are linked to root function, moreover, need to recognize trait variability and the potentially important differences between field- and pot- grown (seedling) roots.