Combined Effects of Partial Defoliation and Nutrient Availability on Cloned Betula pendula Saplings: I CHANGES IN GROWTH, PARTITIONING AND NITROGEN UPTAKE

The combined effects of partial defoliation and nutrient availabilityon dry matter accumulation and partitioning, and on nitrogenuptake and partitioning, were studied in cloned Betula pendulaRoth saplings. The saplings were randomly assigned to differentnutrient levels (5, 1·5 and 0·5 mol Nm^–3)in aerated nutrient culture and to the following defoliationtreatments: (1) control (no damage), (2)damage of the developingmain stem leaves (half of the leaf lamina removed), and (3)removalof the developing main stem leaves (entire leaf lamina removed).Measured in terms of cumulative whole-plant dry weight (includingremoved leaf tissue), the birch saplings were unable to compensatefor the loss of the developing leaves (treatment 3) during the14 d study period. In response to leaf removal (treatment 3)the mean final percentage reduction in whole-plant dry weightwas actually greater than the initial mean percentage reductioncaused by the removal itself; the magnitude of the final reductionwas independent of nutrient availability. After removal of thedeveloping leaves, branch growth was favoured at the expenseof the growth of the rest of the shoot; the relative branchgrowth was most pronounced at the highest nutrient level. Atthe two highest nutrient levels the nitrogen uptake of the saplingswith the developing leaves removed was less than that of undamagedsaplings. We suggest, however, that the incapacity of the saplingsfor compensatory growth after removal of the developing leaveswas primarily due to the decreased total carbon gain of thesaplings rather than to the decreased nitrogen uptake rate. Key words: Partial defoliation, nutrient availability, birch sapling, dry matter, nitrogen     

Combined Effects of Partial Defoliation and Nutrient Availability on Cloned Betula pendula Saplings: II. CHANGES IN NET PHOTOSYNTHESIS AND RELATED BIOCHEMICAL PROPERTIES

The combined effects of partial defoliation and nutrient availabilityon net photosynthesis and related biochemical variables werestudied in cloned Betula pendula Roth saplings. The saplingswere randomly assigned to different nutrient levels (5, 1·5and 0·5 mol N m^–3) in aerated nutrient cultureand to the following defoliation treatments: (1) control (nodamage), (2) damage of the developing main stem leaves (halfof the leaf lamina removed), and (3)removal of the developingmain stem leaves (entire leaf lamina removed). The leaf immediatelybelow the damaged area in the treated plants, and the correspondingleaf in the control plants, were selected for study. Net photosynthesismeasurements and biochemical determinations were made 2, 8 and14 d after assigning the treatments. At intermediate and lownutrient levels the final net photosynthetic capacity was significantlyhigher in the saplings with the topmost leaves removed thanin the undamaged control saplings, indicating that the expressionof compensatory photosynthesis after partial defoliation isnot inhibited by nutrient deficiency. The photosynthetic enhancementwas closely associated with the increased initial activity ofribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco). However,the increased activity of Rubisco was not exclusively the resultof a higher amount of Rubisco. The expression of compensatoryphotosynthesis after partial defoliation in our study cannotunequivocally be attributed to an increased flow of nitrogento the remaining leaves. Key words: Partial defoliation, nutrient availability, net photosynthesis, nitrogen, Rubisco     

Changes in Leaf Gas Exchange Properties of Cloned Betula pendula Saplings after Partial Defoliation

The effects of partial defoliation on the gas exchange characteristicsof the remaining leaves were studied in cloned Betula pendulaL. saplings grown in pots on two different soil types: prefertilizedpeat and unfertilized sand (Experiment 1). The responses ofundamaged leaves to different damage modes of leaf laminae werealso studied using saplings grown on prefertilized peat only(Experiment 2). In Experiment 1, removal of the upper leaves,which represented about half the total leaf area, approximatelydoubled the mean net photosynthetic rates of the remaining lowerleaves on both soil types and at both measuring dates (12–13d and 34 d after assigning the treatments). However, when thelower leaves were removed there was a temporary increase inthe mean net photosynthetic rates of the remaining expandedupper leaves only in the plants grown on sand. In Experiment2, the removal of laminae caused a similar increase in the light-saturatednet photosynthetic rates of the remaining leaves, irrespectiveof whether the laminae were removed totally immediately or graduallyin three stages. The magnitude of the photosynthetic responsewas determined by the amount of leaf tissue removed and wasindependent of the way in which it was removed. We concludethat the increase in the photosynthetic rates of the remainingleaves after partial defoliation may be attributed to the alleviatedcompetitive status among the leaves rather than to the decreasedsource/sink ratio within a plant. Key words: Partial defoliation, net photosynthesis, Betula pendula, birch sapling, herbivory