Light Intensity and Substrate Availability in Relation to Tracheid Development in Picea sitchensis

Assimilation and growth of Picea sitchensis seedlings maintainedat three light intensities were examined in relation to previously-reporteddata on tracheid development in this species. Increases in photosynthetic efficiency with decreased lightlevel failed to compensate completely for reduced total amountsof incident radiation as judged from assimilate distributionpattern and growth rate. Incorporation of assimilate into reservesand new growth declined with decreased light level, while theextent of turnover materials generally increased. Most treatment differences in relative dry weight distributionappeared to result from a combination of differences in growthrate and changes in growth pattern. The proportion of dry weightin branches and roots increased with increased light intensitywhile that in stems and the hypocotyl decreased At the two highest light levels tested, both absolute stem growthrate and total wall accumulation appeared to be independentof substrate availability. This situation contrasts with thatobserved in older material and it is suggested that the initialindependence reflects the temporary existence of a high ratioof functional foliage to stem and cambial material during thefirst year of growth. Since tracheid diameter showed a consistent, positive relationshipwith assimilate status, it is suggested that substrate levelcan exert an important modifying influence on cell expansion.     

Substrate Availability in Relation to Daylength Effects on Tracheid Development in Picea sitchensis (Bong) Carr

Changes in assimilation and growth were studied in Picea sitchensis(Bong) Carr. seedlings initially grown in long days and subsequentlytransferred to short days. These results, and some from a supplementarystudy involving transfer to reduced light quantity alone, wereexamined in relation to tracheid development. Increases in photosynthesis conditioned by reduced daylengthwere greater in magnitude than those resulting from a reductionin light quantity alone. Possible reasons for this as well asthe apparent absence of such photoperiodic effects in the fieldare discussed. Following a reduction in daylength, ¹⁴C incorporation into reservesand new growth generally decreased while turnover materialsincreased in prominence. Some local changes in this generalpattern were apparent, these changes being consistent with growthchanges in the organs under consideration. Initial changes in tracheid wall thickness and diameter aftershort-day transfer appear to be correlated with expected andobserved changes in endogenous substrate, the results for tracheiddiameter confirming a previous observation that substrate levelexerts an important modifying influence on tracheid radial expansion.     

Daylength Effects on Growth, Tracheid Development, and Photosynthesis in Seedlings of Picea sitchensis and Pinus sylvestris

Effects of change in daylength on tracheid expansion and tracheidwall thickening are considered in relation to shoot and rootgrowth, assimilation rate, and starch content, in seedlingsof Picea sitchensis and Pinus sylvestris grown in controlledenviroments. Rates of wall thickening decreased in the first tracheids tobegin wall thickening after decrease in daylength. A sharp decreasein starch-grain size at the same time supports previous suggestionsthat wall thickness is related to available substrate; however,rates of assimilation were found to be far greater in shortdays than in long, suggesting that a surplus of carbohydratemay be produced in long days. It is pointed out that if availablesubstrate does limit wall thickening it is likely to determineto al accumulation of wall material; wall material per tracheidwould follow logistically from this depending on the numberof tracheids around the xylem, and wall thickness would dependalso on the radial tracheid diameter. Unexpectedly, rate of shoot growth accelerated after transferto short days before its final cessation with terminal-bud formation.Root growth declined in short days, but later increased againafter terminal bud formation. Rate of root growth did not appearto be associated with tracheid development in the shoot.     

Analysis of Shoot Apical Growth of Picea sitchensis Seedlings

During the first 100 days after sowing (March-June) the followingchanges took place at the terminal shoot apices of Picea sitchensisseedlings: plastochrones (T) decreased from over 24 h to 4 h;apical domes enlarged from less than 0·20 mm to 0·45mm diameter (D); the ‘projected’ area of tissuesproduced by the apical domes (i.e. viewed from above) increasedin amount from less than 0·012 to 0·024 mm² day^-1;about 15 per cent of this tissue was re-invested in the apicaldomes, the rest was used to produce primordia; and the volume-doublingtimes of the apical dome tissues decreased from over 150 h to50 h. After 100 days there was no further re-investment in theapical domes, but the domes did not decrease abruptly in size.Less tissue was produced per day, but the primordia were smallerso that the rate of primordia formation did not fall precipitously.Plastochrone ratios were inversely related to D, but the relationshipbetween T and D depended on whether T was decreasing or increasing.Progenies which were known to be fast growing tended to build-uptheir apical domes rapidly (i.e. have large ‘re-investmentratios’) and to be capable of producing small primordia.These attributes can evidently be evaluated on seedlings andcould help to lessen the cost of tree breeding progeny-testprogrammes. meristem, Picea sitchensis, Sitka spruce, growth, shoot apex     

Shoot Apical Meristems of Picea sitchensis Seedlings Accelerate in Growth Following Bud-set

First-year seedlings of Picea sitchensis were induced to setbuds by transferring them from long days (17 h) to short days(10 h at 20 °C, 12 h night at 12 °C). After 20 shortdays, the seedlings stopped growing in height, the shoot apicaldomes began enlarging and the transverse growth rates of theshoot apical dome tissues increased. For about 25 days thereafterprimordia were produced faster within the developing buds thanat the apices of seedlings which had been kept in long daysto prevent bud-set. Accelerated primordia production after bud-setenables the largest possible buds to be formed during the frost-freeperiod before winter. Picea sitchensis, apical meristem, bud-set     

Growth response of branches of Picea sitchensis to four years exposure to elevated atmospheric carbon dioxide concentration

Branch bags were used to expose branches on mature Sitka spruce trees to either ambient [CO₂] (A) or elevated [CO₂] (E) for 4 yr. This paper reports the effects of this treatment on the growth, development and phenology of the branches, including shoot expansion, shoot numbers, needle dimensions, needle numbers and stomatal density. The effect of elevated [CO₂] on the relationship between leaf area and sapwood area was investigated. Exposure to elevated [CO₂] doubled photosynthetic rates in current-year shoots and, despite some down-regulation, 1-yr-old E shoots also had higher rates of photosynthesis than their A counterparts. Thus, the amount of assimilate fixed by E branches was substantially more than that fixed by A branches; however, this increase in the local production of assimilate did not lead to an increase in non-structural carbohydrate or stimulate growth or meristematic activity within the E branches. There was a very consistent relationship between leaf area and stem cross-sectional area that was not influenced by [CO₂]. However, unbagged branches had thicker stems than bagged branches, resulting in a slightly lower ratio of leaf area to cross-sectional area. The implications of the results for the modelling of growth and allocation and the potential utility of the branch bag technique are discussed.     

Influence of Environmental Factors on Cell Production and Differentiation in the Early Wood of Picea sitchensis

Three phases are identified in tracheid production by the mainstem of a 12th year forest-grown tree of Picea sitchensis: cellproduction, radial cell expansion and cell wall thickening.For 15 days during formation of early wood daily cell productionwas positively correlated (P=001) with daily solar radiation.Increases in cell production appear related to time-lagged increasesin numbers of cells undergoing expansion and then wall thickening. Radial diameter and wall thickness were measured for each individualcell along the post wall thickened section of radial files oftracheids. The radial diameters of neighbouring cells were positivelycorrelated, groups of 4–5 large cells tending to alternatewith groups of small cells. Furthermore, large cells had thickerwalls than small cells. It is suggested that this distinctivestructure is the result of fluctuations in the environ mentwhich induce changes in the rates of cell production. An increasein cell production increases the length of the queue of cellswaiting to pass successively out of the zones of cell expansionand wall thickening. Consequently cells spend longer in thedevelopment stages, they expand to a greater diameter and theirwalls thicken to a greater thickness than when fewer cells areproduced and the queue is shorter. We suggest that to investigate the influence which weather mayhave on the dynamics of tracheid pro duction it is necessaryto sample developing wood with a frequency twice as great asthat of the frequency of change of the weather variable underconsideration.     

Stilbene glucosides in the bark of Picea sitchensis

The fresh bark of Picea sitchensis contains astringin as the major stilbene; isorhapontin and piceid are present in minor amounts. The aglycones astringenin, isorhaponhgenin and resveratrol are absent in samples of fresh bark. Prenylation reactions on 5,3′,4′-tri-O-methylastringenin are described.     

The Effect of Cold Storage on Subsequent Growth of One-year-old Seedlings of Picea sitchensis

One-year-old seedlings of Picea sitchensis showed a significantdecrease in survival and a decline in vigour in a 5-week growthperiod with periods of cold storage. The effects of cold storagewere most clearly evident in the reduction of main shoot needleweight which would be reflected in the loss of photosyntheticcapacity.     

Shoot Extension in Picea sitchensis II. Analysis of Weather Influences on Daily Growth Rate

The influence of mean temperature and total solar radiationon the daily increment of shoots of 12-year-old Picea sitchensisis described. Serial auto and cross-correlation analysis isused to show that variation in shoot increment is correlatedwith variations in weather occurring over several previous hours.The relationships between daily increments for the leading andtopmost five whorl shoots and the two controlling variablesof temperature and solar radiation are described for the firsthalf of the growing season by auto-regressive models fittedby time-series methods as described by Box and Jenkins. Thesedynamic system models showed that there was a delay of one dayand two days respectively in the influence of temperature andsolar radiation on shoot increment and that after these delaysthe response continued for more than a single day. The maximumfinal response (or steady state gain) of a typical topmost shootto temperature and solar radiation change was found to be 0.091mm d^–1(°C)^–1 and 0.027 mm d^–1 (MJ m^–2)^–1.For the normal range of these variables experienced this indicatesthat shoot extension was five times more sensitive to changesin temperature than to those in solar radiation. Picea sitchensis, Sitka spruce, shoot growth, weather influences, ARMA model, time-series analysis, Box-Jenkins method     

Competition Within Stands of Picea sitchensis and Pinus contorta

Competition was analysed in plots of Picea sitchensis and Pinuscontorta grown in 50 x 50 hexagonal arrays at 14 cm spacingto ages 7 and 5 years, respectively. Relative growth rates in height (RHGR) became positively relatedto tree heights during the year before harvest. Frequency distributionsof tree heights became negatively kurtotic with tendencies towardsleft-skew ness. By the time of harvest, dead trees were evenlydispersed over the plots. Trees with many taller neighbourshad lower RHGRs than trees with few taller neighbours, and theRHGRs of intermediate-sized trees were correlated with their‘competitive status’. Competition was confined mostlyto first-order neighbours. Large trees depressed the RHGRs ofsmaller neighbours and not vice versa; a simple test for this‘one-sided’ competition is described. Neighboursdid not need to greatly overtop a tree to depress its RHGR,they needed only to be at least as tall. Systematic trends inRHGR across the plots, attributed to site heterogeneity, decreasedwith time, and accounted for only about 10 per cent of the variationin RHGR at harvest. Competitive status accounted for 25 and38 per cent of the variation in RHGR in the Picea sitchensisand Pinus contorta plots, respectively. Picea sitchensis, Pinus contorta, competition, monoculture, self-thinning, relative growth rate, growth model     

Adaptive gradients and isolation-by-distance with postglacial migration in Picea sitchensis

Fossil pollen records suggest rapid migration of tree species in response to Quaternary climate warming. Long-distance dispersal and high gene flow would facilitate rapid migration, but would initially homogenize variation among populations. However, contemporary clinal variation in adaptive traits along environmental gradients shown in many tree species suggests that local adaptation can occur during rapid migration over just a few generations in interglacial periods. In this study, we compared growth performance and pollen genetic structure among populations to investigate how populations of Sitka spruce (Picea sitchensis) have responded to local selection along the historical migration route. The results suggest strong adaptive divergence among populations (average Q(ST)=0.61), corresponding to climatic gradients. The population genetic structure, determined by microsatellite markers (R(ST)=0.09; F(ST)=0.11), was higher than previous estimates from less polymorphic genetic markers. The significant correlation between geographic and pollen haplotype genetic (R(ST)) distances (r=0.73, P<0.01) indicates that the current genetic structure has been shaped by isolation-by-distance, and has developed in relatively few generations. This suggests relatively limited gene flow among populations on a recent timescale. Gene flow from neighboring populations may have provided genetic diversity to founder populations during rapid migration in the early stages of range expansion. Increased genetic diversity subsequently enhanced the efficiency of local selection, limiting gene flow primarily to among similar environments and facilitating the evolution of adaptive clinal variation along environmental gradients.     

Effects of Growth Hormone Application on the Secondary Growth of Roots and Stems in Picea sitchensis (Bong.) Carr

Indol-3-ylacetic acid (IAA), gibberellin A3 (GA) and 6 benzylaminopurine(BAP) were applied factorially each at 3x10^–2 M in lanolinto the roots and stems of Sitka spruce seedlings and the activityof the two secondary meristems, the vascular cambium and phellogen,and of the parenchymatous tissues between them, was examined.All the treatments, with the exception of GA produced a localizedstimulation of radial growth at the point of application andthere was a similarity in the response of the various tissuesin both the root and stem. Radial growth of the xylem was notsignificantly affected in the roots whereas in the stems BAPand IAA stimulated growth. In the phloem BAP produced significantstimulation in both roots and stems and IAA stimulated growthin the roots. Growth of the parenchyma and periderm externalto the phloem was also strongly stimulated by both BAP, andIAA in roots and stems. In roots and stems the application of BAP altered the derivativesproduced by the vascular cambium, resulting in the productionof large multiseriate rays in the xylem, and giving rise toan overall increase in the proportion of ray tissue. Picea sitchensis (Bong.) Carr, Sitka spruce, secondary growth, xylem, phloem, periderm, wood rays, Indol-3-ylacetic acid, gibberellin A3, 6 benzylaminopurine, growth hormones     

Photosynthesis and Growth Responses of Parthenocissus quinquefolia (L.) Planch to Soil Water Availability

Photosynthesis and growth characteristics of Parthenocissus quinquefolia were measured under differing soil water availability within a pot. Decreased soil moisture significantly reduced the leaf relative water content (RWC) and the above- and below-ground biomass. However, more biomass was allocated to the root than to the leaf. Net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) were also significantly decreased but water use efficiency (WUE) was increased. Midday depressions in P _N and g _s were not evident for the well-irrigated plants. With the lower water availability, midday reductions in P _N and g _s were much more marked and the duration of the depression was longer. Additionally, the P _N-irradiance response curves also indicated that water supply affected photosynthesis capacity. The growth and photosynthetic response of P. quinquefolia to water supply indicated that this species could resilient to water availabilities and adapt to Hunshandak conditions very well.     

Studies of Growth and Flowering in Picea sitchensis (Bong.) Carr. 1. Effects of Growth Regulator Applications to Mature Scions on Seedling Rootstocks

Effects of growth regulator applications on the flowering of5-years-grafted mature scions of Picea sitchensis (Bong.) Carr.were assessed. Growth regulators were applied to the bud surfacein droplets of ethanol during July, August and September ina polythene house or glasshouse. A mixture of gibberellins A₄and A₇ applied alone, and in combination with gibberellins A₃and A₅, significantly increased numbers by up to seven timesfor male and by up to eight times for female strobili, gibberellinA, gave relatively the strongest response, and gibberellin A₄was inactive. Phosphon D and abscisic acid each reversed thepromotion of flowering by gibberellins, whilst kinetin and N,N-diphenylureahad no effect. The number of female strobili was negativelycorrelated with vegetative shoot length in the year after treatment. Under field conditions hormones were applied in July and Augustunder flaps of bark on the branches of 10-years-grafted maturescions. Gibberellin applications caused a 5-fold increase inflowering and N⁶-benzyladenine further increased the response.Naphth-lyl-acetic acid reduced female and increased male flowering.Bark removal near the base of the branch further enhanced hormone-inducedstrobilus production. The usefulness of these findings for thebreeding of Picea sitchensis is discussed.