Leslie Holdridge defined the subtropical climates as having a mean annual biotemperature between the frost line or critical temperature line, 16 °C to 18 °C (depending on locations in the world), and 24 °C. The frost line separates the warm temperate region from the subtropical region. It represents the dividing line between two major physiological groups of evolved plants. On the warmer side of the line, the majority of the plants are sensitive to low temperatures. They can be killed back by frosts as they have not evolved to withstand periods of cold. On the colder temperate side of the line, the total flora is adapted to survive periods of variable length of low temperatures, whether as seeds in the case of the annuals or as perennial plants that can withstand the cold. The 16 °C–18 °C segment is often "simplified" as 17 °C formula_1.