This review provides a revised and expanded word-formula system of whole-seed primary dormancy classification that integrates the scheme of Nikolaeva with that of Baskin and Baskin. Notable changes include the following. (1) The number of named tiers (layers) in the classification hierarchy is increased from three to seven. (2) Formulae are provided for the known kinds of dormancy. (3) Seven subclasses of class morphological dormancy are designated: ‘dust seeds’ of mycoheterotrophs, holoparasites and autotrophs; diaspores of palms; and seeds with cryptogeal germination are new to the system. (4) Level non-deep physiological dormancy (PD) has been divided into two sublevels, each containing three types, and Type 6 is new to the system. (5) Subclass epicotyl PD with two levels, each with three types, has been added to class PD. (6) Level deep (regular) PD is divided into two types. (7) The simple and complex levels of class morphophysiological dormancy (MPD) have been expanded to 12 subclasses, 24 levels and 16 types. (8) Level non-deep simple epicotyl MPD with four types is added to the system. (9) Level deep simple regular epicotyl MPD is divided into four types. (10) Level deep simple double MPD is divided into two types. (11) Seeds with a water-impermeable seed coat in which the embryo-haustorium grows after germination (Canna) has been added to the class combinational dormancy. The hierarchical division of primary seed dormancy into many distinct categories highlights its great diversity and complexity at the whole-seed level, which can be expressed most accurately by dormancy formulae.

Convallariamajalis has double dormancy and hypogeal germination, but no information is available on embryo growth or on the effects of light and gibberellic acid (GA3) on germination in this genus. Therefore, we investigated embryo growth and other germination features in seeds of C. keiskei and compared the data with those of Trillium camschatcense in another study. Until now, in seeds with double dormancy, embryo growth and germination (epigeal) have been studied in detail only for seeds of T. camschatcense. Phenology of embryo growth and emergence of cotyledonary petiole/root (hereafter root) and shoot in seeds of C. keiskei were monitored outdoors. Effects of temperature, light and GA3 on embryo growth and root and shoot emergence were tested under laboratory conditions. Roots emerged the first spring following seed dispersal in autumn. The embryo grew soon after root emergence, and germination was hypogeal. Seeds with an emerged root formed buds from which a shoot (leaf) emerged above ground during the second spring. Alternating temperatures and light had negative effects on root emergence, and GA3 did not substitute for cold stratification in root emergence. Seeds of C. keiskei have double dormancy, but it differs from that in T. camschatcense. Based on differences in embryo growth before (T. camschatcense) versus after (C. keiskei) root emergence, and on epigeal (T. camschatcense) versus hypogeal (C. keiskei) germination, we suggest that two types of deep simple double morphophysiological dormancy (MPD) be recognized. Since embryo growth in C. keiskei does not fit the standard definition of MPD, we propose to expand this definition.