Effects of soil frost on soil respiration and its radiocarbon signature in a Norway spruce forest soil

Apart from a general increase of mean annual air temperature, climate models predict a regional increase of the frequency and intensity of soil frost with possibly strong effects on C cycling of soils. In this study, we induced mild soil frost (up to −5 °C in a depth of 5 cm below surface) in a Norway spruce forest soil by removing the natural snow cover in the winter of 2005/2006. Soil frost lasted from January to April 2006 and was detected down to 15 cm depth. Soil frost effectively reduced soil respiration in the snow removal plots in comparison to undisturbed control plots. On an annual basis 6.2 t C ha⁻¹ a⁻¹ were emitted in the control plots compared with 5.1 t C ha⁻¹ a⁻¹ in the snow removal plots. Only 14% of this difference was attributed to reduced soil respiration during the soil frost period itself, whereas 63% of this difference originated from differences during the summer of 2006. Radiocarbon (Δ¹⁴C) signature of CO₂ revealed a considerable reduction of heterotrophic respiration on the snow removal plots, only partly compensated for by a slight increase of rhizosphere respiration. Similar CO₂ concentrations in the uppermost mineral horizons of both treatments indicate that differences between the treatments originated from the organic horizons. Extremely low water contents between June and October of 2006 may have inhibited the recovery of the heterotrophic organisms from the frost period, thereby enhancing the differences between the control and snow removal plots. We conclude that soil frost triggered a change in the composition of the microbial community, leading to an increased sensitivity of heterotrophic respiration to summer drought. A CO₂ pulse during thawing, such as described for arable soils several times throughout the literature, with the potential to partly compensate for reduced soil respiration during soil frost, appears to be lacking for this soil. Our results from this experiment indicate that soil frost reduces C emission from forest soils, whereas mild winters may enhance C losses from forest soils.     

Terpenoids are transported in the xylem sap of Norway spruce

Norway spruce is a conifer storing large amounts of terpenoids in resin ducts of various tissues. Parts of the terpenoids stored in needles can be emitted together with de novo synthesized terpenoids. Since previous studies provided hints on xylem transported terpenoids as a third emission source, we tested if terpenoids are transported in xylem sap of Norway spruce. We further aimed at understanding if they might contribute to terpenoid emission from needles. We determined terpenoid content and composition in xylem sap, needles, bark, wood and roots of field grown trees, as well as terpenoid emissions from needles. We found considerable amounts of terpenoids—mainly oxygenated compounds—in xylem sap. The terpenoid concentration in xylem sap was relatively low compared with the content in other tissues, where terpenoids are stored in resin ducts. Importantly, the terpenoid composition in the xylem sap greatly differed from the composition in wood, bark or roots, suggesting that an internal transport of terpenoids takes place at the sites of xylem loading. Four terpenoids were identified in xylem sap and emissions, but not within needle tissue, suggesting that these compounds are likely derived from xylem sap. Our work gives hints that plant internal transport of terpenoids exists within conifers; studies on their functions should be a focus of future research.     

Seasonal dynamics of phloem and xylem formation in silver fir and Norway spruce as affected by drought

The dynamics of phloem growth ring formation in silver fir (Abies alba Mill.) and Norway spruce (Picea abies Karst.) at different sites in Slovenia during the droughty growing season of 2003 was studied. We also determined the timing of cambial activity, xylem and phloem formation, and counted the number of cells in the completed phloem and xylem growth rings. Light microscopy of cross-sections revealed that cambial activity started on the phloem and xylem side simultaneously at all three plots. However, prior to this, 1–2 layers of phloem derivatives near the cambium were differentiated without previous divisions. The structure of the early phloem was similar in silver fir and Norway spruce. Differences in the number of late phloem cells were found among sites. Phloem growth rings were the widest in Norway spruce growing at the lowland site. In all investigated trees, the cambium produced 5–12 times more xylem cells than phloem ones. In addition, the variability in the number of cells in the 2003 growth ring around the stem circumference of the same tree and among different trees was higher on the xylem side than on the phloem side. Phloem formation is presumably less dependent on environmental factors but is more internally driven than xylem formation.     

Effect of local heating and cooling on cambial activity and cell differentiation in the stem of Norway spruce (Picea abies)

BACKGROUND AND AIMS The effect of heating and cooling on cambial activity and cell differentiation in part of the stem of Norway spruce (Picea abies) was investigated. METHODS: A heating experiment (23-25 degrees C) was carried out in spring, before normal reactivation of the cambium, and cooling (9-11 degrees C) at the height of cambial activity in summer. The cambium, xylem and phloem were investigated by means of light- and transmission electron microscopy and UV-microspectrophotometry in tissues sampled from living trees. KEY RESULTS: Localized heating for 10 d initiated cambial divisions on the phloem side and after 20 d also on the xylem side. In a control tree, regular cambial activity started after 30 d. In the heat-treated sample, up to 15 earlywood cells undergoing differentiation were found to be present. The response of the cambium to stem cooling was less pronounced, and no anatomical differences were detected between the control and cool-treated samples after 10 or 20 d. After 30 d, latewood started to form in the sample exposed to cooling. In addition, almost no radially expanding tracheids were observed and the cambium consisted of only five layers of cells. Low temperatures reduced cambial activity, as indicated by the decreased proportion of latewood. On the phloem side, no alterations were observed among cool-treated and non-treated samples. CONCLUSIONS: Heating and cooling can influence cambial activity and cell differentiation in Norway spruce. However, at the ultrastructural and topochemical levels, no changes were observed in the pattern of secondary cell-wall formation and lignification or in lignin structure, respectively.     

Periodicity of cambial activity in Abies balsamea. I. Effects of temperature and photoperiod on cambial dormancy and frost hardiness

The relationship between from hardiness and growth potential, and their dependence on temperature and photoperiod, was investigated in the one-year-old cambium of balsam fir [Abies balsamea (L.) Mill.]. Six-year-old trees were exposed for 9 weeks to either the natural environment or one of 4 controlled environments in the fall (18 September-18 November), spring (12 April–14 June) and summer (19 July – 19 September). The 4 controlled environments were (1) WS, warm temperature (24/20°C in day/night) + short day (8 h). (2) WL. warm temperature (24/20°C) + long day (8 h + 1 h night break), (3) CS. cold temperature (9/5°C) + short day (8 h) and (4) CL, cold temperature (9/5°C) + long day (8 h + 1 h night break). At the beginning and end of each exposure, cambial activity was measured by recording the number of xylem, cambium and phloem cells, frost hardiness was estimated from the cambium's ability to survive freezing to –40°C, and cambial growth potential was deduced from the duration of the cell cycle and the production of xylem, cambium and phloem cells in cuttings cultured for 4 weeks with exogenous indole-3-acetic acid (IAA) under environmental conditions favourable for cambial activity. In the natural environment, frost hardening began in September and was completed in November, while dehardening occurred when the cambium reactivated. CL, CS, and to a lesser extent WS, promoted hardening in the summer and fall, but did not prevent dehardening in the spring. The cambial growth potential in the natural environment declined from a maximum in April to a low level in June, reached a minimum in September, then increased to a high level in November. This potential was promoted by CL and CS on all dates by WL in the summer and fall. The ratio of xylem to phloem induced by IAA treatment was greatest in June and least in September in cuttings from trees exposed to the natural environment, and was increased by CL and CS in the fall. The cambium in intact branches of trees protected from chilling during the fall and winter resumed cell cycling after less than 9 weeks of dormancy, but produced mostly or only phloem in the subsequent growing period. It is concluded that the frost hardiness of the cambium, the IAA-induced cycling of cambial cells, and IAA-induced xylem to phloem ratio vary independently with season, temperature and photoperiod, and that the periodicity of these processes is regulated endogenously.     

Ethylene effects on cambial activity and cell wall formation in hypocotyls of Picea abies seedlings

Ethylene regulation of cell division in the vascular cambium and cell wall formation was studied in hypocotyls of Norway spruce ( Picea abies [L.] Karst.) seedlings. Cuttings from 6-week-old seedlings were placed in water culture to which compounds affecting the synthesis and action of ethylene were added. After a 3-week treatment period, growth, ethylene production, morphology and cell wall composition of the hypocotyls were determined. Addition of high concentrations of the potent ethylene releasing agent 2-chloroethylphosphonic acid (ethrel), which increased ethylene emission by more than twice compared to control plants, inhibited the expansion of xylem cells while stimulating the incorporation of cell wall material, especially cellulose. Addition of small amounts of ethrel, which slightly stimulated ethylene emission, led to increases in the size of xylem cells, the amount of phloem tissue and the number of intercellular spaces in the cortex, and thus to increased hypocotyl diameter. However, no significant change in cell wall composition was detected. When ethylene production was decreased by adding Co²⁺ to the nutrient solution, differentiation of new xylem was disturbed, but the rate of cell division was not affected. Although the incorporation of cell wall material was inhibited, the proportions of lignin and cellulose in the wall appeared to remain unchanged. Silver ions stimulated the expansion of both xylem and cortex cells, but had no significant effect on cell wall formation. We conclude that ethylene has a role in regulating the incorporation of wall carbohydrates.     

Wood diameter indicates diurnal and long-term patterns of xylem water potential in Norway spruce

The stem diameter of adult Norway spruce trees was measured to see whether changes in xylem water potential lead to detectable radial deformation of the wood. The dendrometers used in these experiments measured only the dimensional changes of the woody cylinder (sap- and heartwood). Wood diameter was measured close to the ground and just below the living crown. After correction for thermal expansion of dendrometers and wood, diurnal variation of wood diameter ranged between 50 and 180 µm. Psychrometric measurements showed that xylem water potential varied in parallel to wood diameter. Diameter changes were always more pronounced at the higher stem position and exhibited a clear diurnal pattern. During the day, wood diameter decreased with increasing vapor pressure deficit and transpiration rate and with decreasing twig water potential. At night, the wood re-expanded but did not always reach the dimension of the previous day. Pre-dawn wood diameter decreased during periods of soil drought, a process which rapidly stopped and reversed after rain events. On several days, oscillation in wood diameter was observed during the mid-day hours. The oscillation had a period of approximately 50 min and showed a phase shift between different stem heights. All observed patterns of wood shrinkage and expansion were consistent with the hypothesis that xylem water tension leads to an elastic contraction of xylem conduits. The results demonstrate that xylem diameter is more suitable than whole-stem diameter for monitoring changes in xylem water potential.     

Predicting spring frost sensitivity by bud development and temperature sum in Norway spruce seedlings

Nurseries would benefit greatly if frost hardiness (FH) of seedlings could be predicted by some environmental variable or by bud development in spring. We investigated the FH of 1-year-old Norway spruce (Picea abies (L.) Karst.) seedlings of local origin. The seedlings were stored frozen until incubated in the growth chamber at six different temperature sums (TSs) (0, 55, 88, 142, 185 and 240 d.d., >5°C) from mid-February to mid-March. FH of the buds, stems and previous year needles was assessed on three occasions. When the TS was 88 d.d. or less, buds exhibited only microscopic signs of development, even when seedlings tolerated temperatures below −10°C. As TS increased, primordial needles and primordial stems of buds grew while FH weakened, especially in previous year needles. When the TS was at least 142 d.d., all plant parts were frost hardy to approximately −6°C. Monitoring TS and bud development can help predict FH of Norway spruce seedlings in spring. However, more studies with seedlings of different ages and from multiple locations are necessary to appreciate the generality of our results.     

Short-term effects of nutrient compensation following whole-tree harvesting on soil and soil water chemistry in a young Norway spruce stand

A growing demand for bioenergy from conventional forestry in Sweden will increase the need of nutrient compensation, that preferably should be made relatively shortly after harvesting and have no undesired side-effects. This study compared the effects of granulated wood ash (Ash), N-free, dolomite-based fertiliser (Vitality) and the green fraction of harvest residues (Residues) on the podsolic soil and soil solution of a young Norway spruce (Picea abies (L.) Karst) stand in SW Sweden. The treatments were applied three years after clear-felling and whole-tree harvesting. The soil solution was repeatedly sampled in the rooting zone 2–5 years after treatment. The soil study was performed 4 years after the Ash treatment and 3 years after Residues treatment and the last Vitality treatment (the Vitality treatment was applied on two occasions over 2 years). The Vitality treatment increased base saturation and effective CEC in the humus layer in relation to the other treatments, and also increased Ca and K concentrations in the soil solution. The Ash treatment resulted in higher exchangeable K concentration than Vitality in the litter layer, and Residues increased K concentrations in the soil water. No treatment influenced the KCl-exchangeable nitrate concentrations in the soil or the nitrate levels in the soil water. The results indicate that granulated wood ash could be used for long-term nutrient compensation without undesired short-term side-effects.     

Some effects of long-term ozone fumigation on Norway spruce

Summary Potted cuttings of a 12-year-old Picea abies tree were fumigated with ozone, 100 or 300 g O₃· m^–3 (50 or 150 ppb O₃) being added to charcoal-filtered air during the 1985 growing season for a total of 1215 h. The wax structure of ozone-fumigated needles was no different from that of controls. Because flattened wax structures and fused wax fibrils also occurred in controls, these phenomena could not serve as bioindications for the ozone concentrations applied. A smooth layer was found beneath the soluble wax layer and covered needle surface and stomatal openings of ozone-fumigated needles to a greater extent than in controls. Wax quantity was considerably reduced by fumigation with 300 g O₃ · m^–3. Leaf pigments (as extracted with the wax) were less abundant in needles treated with 300 gO₃; the smooth layer probably contributed to the impeded extraction of pigments.     

Some effects of long-term ozone fumigations on Norway spruce

Summary Potted cuttings of a 12-year-old, and grafts of an 80-year-old, Norway spruce (Picea abies (L.) Karst.) were subjected to 100 or 300 g O₃·m^–3 for 1215 h (45 h of daylight per week) during the growing season of 1985. At the end of the fumigation the plants did not exhibit any visible signs of injury. Whereas in the fumigation with 100g O₃·m^–3 we did not detect any significant change in gas exchange, 300 g O₃·m^–3 did alter the CO₂ uptake after 27 weeks, and in one clone transpiration was also altered. Stomatal reaction to a change of light suggested sluggishness, but the change was not statistically significant.     

The elemental composition, the microfibril angle distribution and the shape of the cell cross-section in Norway spruce xylem

Relationships between the elemental composition, the microfibril angle (MFA) distribution and the average shape of the cell cross-section of irrigated-fertilised and untreated Norway spruce (Picea abies [L.] Karst.) earlywood were studied. Sample material was obtained from Flakaliden, Sweden. The elemental composition was studied by determining the relative mass fractions of the elements P, S, Cl, K, Ca and Mn by X-ray fluorescence and by determining the mass absorption coefficients for X-rays. X-ray diffraction was used to determine the MFA distribution and the average shape of the cell cross-section. The latter was also determined by light microscopy. In transition from juvenile wood to mature wood, a decrease of the mode of the MFA distribution from 13°–24° to 3°–6° was connected to a change in the shape of the cell cross-section from circular to rectangular. The irrigation-fertilisation treatment caused no change in the MFA distribution or in the shape of the cell cross-section, whereas the mass absorption coefficient was higher and the density was smaller in irrigated-fertilised wood. Larger proportion of the elements S, Cl and K, but smaller proportion of the element Mn, were observed due to the treatment. The results indicate that the shape of the cell cross-section or the MFA distribution are not directly linked to the growth rate of tracheids or to the nutrient-element content in the xylem and only show notable changes as a function of the cambial age.     

Intra-annual tracheid formation of Norway spruce provenances in southern Finland

We studied the intra-annual wood formation in a Norway spruce provenance experiment in southern Finland from 2004–2008. Two Finnish provenances, northern and southern, as well as German and Hungarian provenances were included. Timing of tracheid formation and differentiation, and tracheid dimensions were determined from periodically extracted microcores. The aim was to determine the differences between the years and provenances in the timing of the xylogenesis and in the xylem characteristics. Year-to-year variation was high both in timing of tracheid formation and xylem characteristics, while between-provenance differences were small. The onset of tracheid formation varied from early May to late June in different trees in different years. The onset of tracheid formation was not closely related to the annual variations of temperature sum. In all the years, daily temperatures exceeded the threshold +5°C for several weeks before the onset of tracheid formation. The highest tracheid formation rate occurred after the summer solstice in all years and generally coincided with the highest daily temperatures during the growing season. Tracheid production ceased early in 2006 due to a mid-summer drought. Cell differentiation continued late in autumn as non-mature tracheids were still observed around mid-September. No clear differences between the provenances in the timing of tracheid formation were observed, although the Finnish provenances tended to initiate tracheid formation slightly earlier than the other provenances. The tree-ring widths of the Finnish provenances were also wider, while tracheid diameter of the German provenance was slightly smaller. Our results indicate that between-tree variation in the timing of wood formation is high compared with the latitude effect of seed source.     

The effects of enhanced UV-B radiation on physiological activity and growth of Norway spruce planted outdoors over 5 years

The responses of Norway spruce [Picea abies (L.) Karst.] to enhanced UV-B radiation during the 5-year treatment performed outdoors have been subjected to ecophysiological and growth analysis. The plants were exposed to UV-B radiation, simulating 17% ozone depletion. Ecophysiological parameters were monitored three times a year on three needle age classes, while growth was analysed at the end of each growth season. Spruce exhibited great variability in the amounts of photosynthetic pigments and methanol-soluble UV-B absorbing compounds, light use efficiency, photosynthesis and respiratory potential. The needle, branch and plant biomass production was not significantly affected during the 5-year treatment. The repeated-measures procedure comparing growth parameters through subsequent seasons, revealed a decrease of branch diameter under enhanced UV-B, which could be interpreted as a cumulative UV-B effect. The effects of UV-B radiation depended on needle development stage, interaction with environmental conditions and stresses. A reduced negative effect of UV-B radiation was observed during the prolonged drought in 2003, which was hypothesised as an alleviating effect. The tolerance of Norway spruce to elevated UV-B was to a large extent due to the high content of methanol-soluble UV-B absorbing compounds that was related neither to environmental conditions, including UV-B dose, nor to the developmental stage of the needles. The current year needles exhibited a tendency to increased production of UV-B absorbing compounds under elevated UV-B radiation. The outdoor study performed under variable environmental conditions showed great complexity of spruce response to enhanced UV-B.     

The use of different soil nitrogen sources by young Norway spruce plants

 Three-year-old Norway spruce trees were planted into a low-nitrogen mineral forest soil and supplied either with two different levels of mineral nitrogen (NH₄NO₃) or with a slow-release form of organic nitrogen (keratin). Supply of mineral nitrogen increased the concentrations of ammonium and nitrate in the soil solution and in CaCl₂-extracts of the rhizosphere and bulk soil. In the soil solution, in all treatments nitrate concentrations were higher than ammonium concentrations, while in the soil extracts ammonium concentrations were often higher than nitrate concentrations. After 7 months of growth, ¹⁵N labelled ammonium or nitrate was added to the soil. Plants were harvested 2 weeks later. Keratin supply to the soil did not affect growth and nitrogen accumulation of the trees. In contrast, supply of mineral nitrogen increased shoot growth and increased the ratio of above-ground to below-ground growth. The proportion of needle biomass to total above-ground biomass was not increased by mineral N supply. The atom-% ¹⁵N was higher in younger needles than in older needles, and in younger needles higher in plants supplied with ¹⁵N-nitrate than in plants supplied with ¹⁵N-ammonium. The present data show that young Norway spruce plants take up nitrate even under conditions of high plant internal N levels. Received: 1 April 1998 / Accepted: 9 October 1998     

The activity of uridine diphosphate-D-glucose: Nicotinamide-adenine dinucleotide oxidoreductase in cambial tissue and differentiating xylem isolated from sycamore trees

Summary The activity of UDPGlc: NAD oxidoreductase is measured in enzyme preparations obtained from sycamore cambium and xylem tissue. The activity of this enzyme is greater in xylem than in cambium whether expressed on a specific activity basis or on a per-cell basis. It is suggested that, in developing xylem, direct oxidation of UDPGlc may contribute significantly to the biosynthesis of polysaccharide precursors.Abbreviations UDPGlc Uridine-Diphosphate-D-Glucose - UDPGlcA Uridine Diphosphate-D-Glucuronic Acid - UDPXyl Uridine Diphosphate-D-Xylose - NAD Nicotinamide-Adenine dinucleotide - D-GlcA D-Glucuronic acid - E.C. Enzyme Commission - Ci Curie     

Nitrogen transformations in limed and nitrogen fertilized soil in Norway spruce stands

Nitrogen transformations in the soil, and the resulting changes in carbon and nitrogen compounds in soil percolate water, were studied in two stands of Norway spruce (Picea abies L.). Over the last 30 years the stands were repeatedly limed (total 6000 kg ha^–1), fertilized with nitrogen (total about 900 kg ha^–1), or both treatments together. Both aerobic incubations of soil samples in the laboratory, and intact soil core incubations in the field showed that in control plots ammonification widely predominated over nitrification. In both experiments nitrogen addition increased the formation of mineral-N. In one experiment separate lime and nitrogen treatments increased nitrification, in the other, only lime and nitrogen addition together had this effect. In one experiment immobilization of nitrogen to soil microbial biomass was lower in soil only treated with nitrogen. Soil percolate water was collected by means of lysimeters placed under the humus layer and 10 cm below in the mineral soil. Total N, NH₄-N and NO₃-N were measured, and dissolved organic nitrogen was fractioned according to molecular weight. NO₃-N concentrations in percolate water, collected under the humus layer, were higher in plots treated with N-fertilizer, especially when lime was also added. The treatments had no effect on the N concentrations in mineral soil. A considerable proportion of nitrogen was leached in organic form.     

The dibenzodioxocin lignin substructure is abundant in the inner part of the secondary wall in Norway spruce and silver birch xylem

A specific condensed lignin substructure, dibenzodioxocin, was immunolocalized in differentiating cell walls of Norway spruce (Picea abies (L.) H. Karsten) and silver birch (Betula pendula Roth) xylem. A fluorescent probe, Alexa 488 was used as a marker on the dibenzodioxocin-specific secondary antibody. For the detection of this lignin substructure, 25-m cross-sections of xylem were viewed with a confocal laser-scanning microscope with fluorescein isothiocyanate fluorescence filters. In mature cells, fluorescence was detected in the S3 layer of the secondary wall in both tree species, but it was more intense in Norway spruce than in silver birch. In silver birch most of the signal was detected in vessel walls and less in fiber cell walls. In very young tracheids of Norway spruce and vessels and fibers of silver birch, where secondary cell wall layers were not yet formed, the presence of the dibenzodioxocin structure could not be shown.Abbreviation CLSM confocal laser-scanning fluorescence microscopy     

Differences in frost hardiness of two Norway spruce morphotypes growing at Mt. Brocken, Germany

Norway spruce (Picea abies (L.) Karst.) exhibits strong ecotypic variation along altitudinal gradients in morphological traits, e.g. slenderness of crowns or arrangement of second-order branches. We were interested whether montane and lowland morphotypes differ in a key trait for the survival in cold environments, i.e. frost hardiness, and asked: (i) are montane morphotypes more resistant to frost damage and (ii) do they have a lower risk of frost damage by late frosts in spring than lowland morphotypes?We used the electrolyte leakage-method to measure frost hardiness on a monthly basis from October 2006 to May 2007 in stands of the montane and lowland morphotypes at Mt. Brocken in the Harz Mountains, Germany.LT₅₀ (i.e. the temperature that results in 50% of maximum electrolyte leakage) was assessed by freezing treatments in a frost chamber and was significantly influenced by morphotype, month and minimum ambient temperatures. LT₅₀ was significantly lower in the montane than in the lowland morphotype, with −107 °C and −49 °C, respectively. However, the interactions between morphotype with minimum ambient temperature or month were not significant. Thus, as frost hardiness of the two morphotypes responded to temperature in the same way, both morphotypes can be supposed to be exposed to the same risk of frost damage during hardening in autumn and dehardening in spring.     

The regulation of cambial division and secondary xylem differentiation in xanthium by auxins and gibberellin

Cambial division continued in decapitated Xanthium plants without concomitant xylem fiber differentiation. The application of indoleacetic acid to these plants did not affect the production of cambial derivatives or induce xylem fiber differentiation. When naphthaleneacetic acid was applied either to the second internode or to the stump of a lateral shoot, xylem fiber differentiation was induced in the newly formed cambial derivatives on the xylem side of the cambium in the stem. When naphthaleneacetic acid was applied unilaterally, xylem fiber differentiation was restricted to that side of the stem in the first internode and hypocotyl. Naphthaleneacetic acid also enhanced the production of cambial derivatives. Gibberellic acid enhanced cambial derivative production but did not affect the differentiation of xylem fibers. Similar numbers of cambial derivatives were produced in some naphthaleneacetic acid-treated plants in which xylem fiber differentiation was induced and in gibberellic acid-treated plants which did not differentiate xylem. When naphthaleneacetic acid was applied 72 hours after decapitation, the oldest of the cambial derivatives on the xylem side failed to develop into fibers although younger cells did. These results suggest that auxin has its direct effect on the induction of xylem differentiation rather than the induction of divisions prerequisite to differentiation.