Abstract:Understanding the relationships between twig and leaf traits provides valuable insights into the adaptive strategies of plant species. There are limited studies addressing the shade tolerance of woody species regarding trade-offs between twig and leaf traits. However, there are two contrasting hypotheses regarding the characteristics of shade tolerance; whether shade-tolerant species prefer maximized light interception and low respiration in the shade or are characterized by tough structures because of a reduction in organ turnover and promotion of plant survival. In this study, our objectives were to reveal whether the twigs and leaves of shrubs exerted trade-offs in the shade. Thus, twig dry weight (TDW), leaf dry weight unit twig (LDW), leafing intensity (LN/TDW), specific twig length (TL/TDW), leaf area supporting efficiency (LA/TDW), and individual leaf area (ILA) were measured for 11 shrub species, which have a relatively high ornamental value and limited management under the canopy at the Beijing Forestry University campus. In addition, the relative light summation of photosynthetically active radiation (PAR) and spectral ratio of red light and far-red light (R/FR) were measured for the habitats of target shrubs. Relationships between twig and leaf traits were determined using standardized major axis estimation (SMA). Effects of shade on relationships between twig and leaf traits were estimated using a multiple regression analysis. The study showed that (1) light (relative light summation of PAR and R/FR) has significant positive relationships (P < 0.01) with TDW and LDW and significant negative relationships (P < 0.001) with TL/TDW, LA/TDW, and LN/TDW, whereas no significant relationship (P > 0.05) was observed with ILA. In addition, the interactions between light and traits were more significant for R/FR than the relative light summation for PAR; (2) furthermore, there was a significant positive correlation (P < 0.001) and allometric relationship between TDW and LDW. A direct relation was observed between light and biomass of twigs and leaves, as biomass of both significantly decreased with decreasing light. However, shrubs reflected a relative bias in investment of leaf biomass; (3) LA/TDW also showed significant (P < 0.001) positive correlations and isometric relationships with TL/TDW. However, the steepness for TL/TDW-LA/TDW relationships in the lower R/FR decreased, indicating that shrubs presented a relatively conservative strategy for light interception in the shade; (4) ILA showed a significant (P < 0.001) negative correlation and allometric relationship with LN/TDW, implying that the twigs of shrubs tended to be made up of many smaller leaves in the shade. Therefore, together, the trade-offs of ornamental shrubs were similar to the hypothesis that suggested shade-tolerant species exert trade-offs in growth performance. However, generally, the shade tolerance of these shrubs was too weak to allow them to grow in the shade as ornamental species.