Warming and elevated ozone differently modify needle anatomy of Norway spruce (Picea abies) and Scots pine (Pinus sylvestris)

Acclimation of conifer needle anatomy to climate change is poorly understood. We studied needle anatomy, shoot gas exchange, current-year shoot length, and stem diameter growth in Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) seedlings exposed to elevated ozone (1.35× to 1.5× ambient concentration) and elevated temperature (0.9–1.3 °C + ambient temperature) alone and in combination for two exposure seasons in two separate open-field experiments in central Finland. Pines grew also at two soil nitrogen levels. In spruce, warming increased mesophyll intercellular space and reduced gas exchange and shoot growth and made needles narrower and the epidermis and hypodermis thinner. In pine, warming made needles bigger, increased shoot and stem growth, stomatal row number, and proportions of vascular cylinder, phloem, and xylem and reduced the proportion of mesophyll. These responses indicate that pine benefited and spruce suffered from moderate warming. Ozone caused a thickening of epi- and hypo-dermis and a lower stomatal conductance in both species, reduced stomatal density in spruce, and increased proportions of phloem, xylem, and sclerenchyma and reduced growth in pine. Ozone responses suggest increased oxidative stress defense. Stomatal responses were affected by interactions of elevated temperature and ozone in both species. Nitrogen availability modified ozone and temperature responses, particularly in the vascular tissues in pine.