They also had larger crowns, suggesting that dense stems provide strength for supporting larger crowns. Shade-tolerant species had denser wood and slower extension growth. Specific leaf area (SLA, leaf area/leaf mass) and wood density were the traits most strongly correlated to shade tolerance and extension growth. Photosynthetic capacity, leaf dark respiration and total leaf mass related weakly to shade tolerance and extension growth. Plant traits that related positively with shade tolerance were negatively related to extension growth, and vice versa. Shade tolerance and extension growth of the leader shoot were negatively correlated. Growth was determined from annual lengths of leader shoots, and shade tolerance from an independently tested scale. Plant traits were measured for 0.5-1 m tall saplings: 10 saplings growing at low light conditions and 10 at high light conditions. To explore how such differences depend on underlying plant traits, 14 tree species were investigated in temperate forests on sand and loess soils in the Netherlands. Species differences in growth and shade tolerance might contribute to coexistence of tree species.
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