European Larch and Eastern White Pine Respond Similarly During Three Years of Partial Defoliation

To test whether trees with different leaf life spans respond differently to defoliation, eastern white pine (Pinus strobus L.) and European larch (Larix decidua Mill.) trees (9 years old in 1991) were partially defoliated by hand between July 1 and 10 in 1989, 1990 and 1991. At the end of 1991, trees of both species had received either 0, 1, 2 or 3 years of defoliation. Trees that received only 1 year of defoliation were defoliated in 1989. Variables measured included photosynthesis, twig water potential, leaf mass per area and leaf nitrogen concentration. There were few significant responses to defoliation in any of the three years of treatment in either species, and only the current-year defoliation treatments caused significant responses. Both species had reduced photosynthetic rates and less negative twig water potentials in response to defoliation in 1989. In 1990 and 1991, the defoliation treatments had no significant effect on any of the parameters measured in European larch. In 1990, there was a significant reduction in foliar nitrogen concentration in eastern white pine in response to defoliation in 1990. In 1991, eastern white pine had significantly less negative twig water potentials in response to defoliation in 1991. Leaf mass per area was not affected by defoliation in either species. We conclude that, for European larch and eastern white pine, differences in leaf life span have no effect on leaf-and twig-level responses to defoliation

The Effect of Defoliation Intensity and History on Photosynthesis, Growth and Carbon Reserves of Two Conifers with Contrasting Leaf Lifespans and Growth Habits

The effects of partial defoliation on photosynthesis, whole-seedling carbon allocation, partitioning and growth were studied for two species with contrasting foliar traits. Field-grown seedlings of deciduous Japanese larch (Larix leptolepis) and evergreen red pine (Pinus resinosa) were defoliated by hand in early summer for 2 consecutive years. In the first year (1990), seedlings were defoliated by removing the distal 0, 25, 50 or 75% of each needle. In the second year (1991), seedlings were defoliated either 0 or 50%, regardless of previous defoliation treatments. Defoliation had little effect on photosynthesis and starch concentration in whole seedlings of either species in the first year. In the second year, photosynthesis increased in both species in response to the 1991 defoliation treatment, and in red pine also increased in response to the 1990 defoliation treatment. Further, in 1991 both larch and pine had decreased whole-seedling total non-structural carbohydrate concentrations in all seedlings that were defoliated at least once over the 2-yr period. This decrease was noted mostly in the starch component of the nonstructural carbohydrates, and was similar in both species. In 1991, biomass was similarly decreased in both species in response to 1991 defoliation. Both species showed overcompensation in total and component biomass in seedlings defoliated by 25% in 1990. Overall, the results do not support the widely held belief that evergreen trees are substantially more affected than deciduous trees by defoliation

Effects of Elevated Ozone and Low Light on Diurnal and Seasonal Carbon Gain in Sugar Maple

The long-term interactive effects of ozone and light on whole-tree carbon balance of sugar maple (Acer saccharum Marsh.) seedlings were examined, with an emphasis on carbon acquisition, foliar partitioning into starch and soluble sugars, and allocation to growth. Sugar maple seedlings were fumigated with ambient, 1·7 × ambient and 3·0 × ambient ozone in open-top chambers for 3 years under low and high light (15 and 35% full sunlight, respectively). Three years of ozone fumigation reduced the total biomass of seedlings in the low- and high-light treatments by 64 and 41%, respectively, but had no effect on whole-plant biomass allocation. Ozone had no effect on net photosynthesis until late in the growing season, with low-light seedlings generally exhibiting more pronounced reductions in photosynthesis. The late-season reduction in photosynthesis was not due to impaired stomatal function, but was associated more with accelerated senescence or senescence-like injury. In contrast, the 3·0 × ambient ozone treatment immediately reduced diurnal starch accumulation in leaves by over 50% and increased partitioning of total non-structural carbohydrates into soluble sugars, suggesting that injury repair processes may be maintaining photosynthesis in late spring and early summer at the expense of storage carbon. The results in the present study indicate that changes in leaf-level photosynthesis may not accurately predict the growth response of sugar maple to ozone in different light environments. The larger reduction in seedling growth under low-light conditions suggests that seedlings in gap or closed-canopy environments are more susceptible to ozone than those in a clearing. Similarly, understanding the effects of tropospheric ozone on net carbon gain of a mature tree will require scaling of leaf-level responses to heterogeneous light environments, where some leaves may be more susceptible than others

Effects of Elevated Ozone on CO2 Uptake and Leaf Structure in Sugar Maple Under Two Light Environments

The interactive effects of ozone and light on leaf structure, carbon dioxide uptake and short-term carbon allocation of sugar maple (Acer saccharum Marsh.) seedlings were examined using gas exchange measurements and 14C-macroautoradiographic techniques. Two-year-old sugar maple seedlings were fumigated from budbreak for 5 months with ambient or 3 x ambient ozone in open-top chambers, receiving either 35% (high light) or 15% (low light) of full sunlight. Ozone accelerated leaf senescence, and reduced net photosynthesis, 14CO2 uptake and stomatal conductance, with the effects being most pronounced under low light. The proportion of intercellular space increased in leaves of seedlings grown under elevated ozone and low light, possibly enhancing the susceptibility of mesophyll cells to ozone by increasing the cumulative dose per mesophyll cell. Indeed, damage to spongy mesophyll cells in the elevated ozone x low light treatment was especially frequent. 14C macroautoradioraphy revealed heterogeneous uptake of 14CO2 in well defined areole regions, suggesting patchy stomatal behaviour in all treatments. However, in seedlings grown under elevated ozone and low light, the highest 14CO2 uptake occurred along larger veins, while interveinal regions exhibited little or no uptake. Although visible symptoms of ozone injury were not apparent in these seedlings, the cellular damage, reduced photosynthetic rates and reduced whole-leaf chlorophyll levels corroborate the visual scaling of whole-plant senescence, suggesting that the ozone x low light treatment accelerated senescence or senescence-like injury in sugar maple

Response of Japanese Barberry to Varying Degrees of Defoliation

Until recently, it was thought that Berberis thunbergii (Japanese Barberry), a non-native invasive plant that has become particularly widespread in certain regions of New Jersey, benefited from a lack of herbivorous defoliators. However, in 2007 extensive defoliation was documented across a wide geographical distribution in New Jersey, calling this assumption into question. We tested whether Japanese Barberry was negatively affected by partial defoliation by manually clipping 50% or 100% of leaves on current-year stems on small and large plants in the summer of 2008. We found almost no impact of defoliation on growth, carbon storage, or leaf-level physiology for either treatment. We noted some differences between large and small plants, but these were not related to defoliation treatments. Our results suggest that, even in the presence of herbivory, Japanese Barberry is capable of maintaining growth and carbon reserves, thus making it an effective competitor for resources

White Pine, Japanese Larch, and Bear Oak Respond Differently to Partial Defoliation

Seedlings (2-0 bare root) of Japanese larch (Larix leptolepis), white pine (Finns strobus), and bear oak (Quercus ilicifolia) were partially defoliated by clipping half of each leaf per plant. Control plants were not clipped. Photosynthesis, water potential, leaf nitrogen concentration, leaf mass, and total mass were measured one week and nine weeks after treatment. Compensatory responses varied by species and timing. Defoliated Japanese larch had increased photosynthesis rates and foliar nitrogen concentration relative to control plants one week after treatment. Total biomass was also increased relative to controls by the end of the experiment. Defoliated pine seedlings had increased leaf mass and total biomass relative to controls by the end of the experiment. Defoliated oak seedlings showed no compensatory responses. Leaf lifespan did not appear to be a determinant of response amount or type. Instead, we suggest that degree of allocation of resources to aboveground growth, as indicated by root-to-shoot ratio, may be related to degree of compensation to partial defoliation

Plant Size, Not Age, Regulates Growth and Gas Exchange in Grafted Scots Pine Trees

We studied the effect of scion donor-tree age on the physiology and growth of 6- to 7-year-old grafted Scots pine (Pinus sylvestris L.) trees (4 and 5 years after grafting). Physiological measurements included photosynthethetic rate, stomatal conductance, transpiration, whole plant hydraulic conductance, needle nitrogen concentration and carbon isotope composition. Growth measurements included total and component biomasses, relative growth rates and growth efficiency. Scion donor trees ranged in age from 36 to 269 years at the time of grafting. Hydraulic conductance was measured gravimetrically, applying the Ohm\u27s law analogy, and directly, with a high-pressure flow meter. We found no effect of scion donor-tree age on any of the variables measured. There was, however, great variation within scion donor-tree age groups, which was related to the size of the grafted trees. Differences in size may have been caused by variable initial grafting success, but there was no indication that grafting success and age were related. At the stem level, hydraulic conductance scaled with total leaf area so that total conductance per unit leaf area did not vary with crown size. However, leaf specific hydraulic conduc-tance (gravimetric), transpiration, photosynthesis and stomatal conductance declined with increasing total tree leaf area and needle width. We hypothesize that needle width is inversely re-lated to mesophyll conductance. We conclude that canopy and needle size and not scion donor-tree age determined gas ex-change in our grafted trees

Tree Height and Age-Related Decline in Growth in Scots Pine (Pinus Sylvestris L.)

Growth and seasonal water use was measured amongst trees growing in an old growth Scots pine forest in the Scottish Highlands. Three sites which differed in their recent management history and contained old and naturally regenerated young trees growing together were monitored in the field. Our results showed a clear decrease in growth efficiency with age, from values of around 0.25 kg m-2 leaves year-1 in approximately 25-year-old trees to less than 0.1 kg m-2 leaves year-1 in trees over 200 years old. When the old trees in one of the field sites were released from competition by thinning, their growth efficiency reverted to that of coexisting young trees, indicating that the decline in growth was reversible. This is consistent with the results of a parallel study showing that cambial age had no effect on the physiology or growth of grafted seedlings originating from the same population studied here (Mencuccini et al. 2005). Our detailed study of tree water use in the field showed an overall decrease in whole-tree hydraulic conductance and stomatal canopy conductance with tree height in the unthinned stands, in agreement with the hydraulic limitation hypothesis. However, the effect of this reduction in hydraulic efficiency on growth was comparatively small, and old trees also showed consistently lower nitrogen concentrations in needles, suggesting that hydraulic and nutritional factors combined to produce the decline in growth efficiency with age observed in the studied populations