Shade, light, and stream temperature responses to riparian thinning in second-growth redwood forests of northern California.

Resource managers in the Pacific Northwest (USA) actively thin second-growth forests to accelerate the development of late-successional conditions and seek to expand these restoration thinning treatments into riparian zones. Riparian forest thinning, however, may impact stream temperatures-a key water quality parameter often regulated to protect stream habitat and aquatic organisms. To better understand the effects of riparian thinning on shade, light, and stream temperature, we employed a manipulative field experiment following a replicated Before-After-Control-Impact (BACI) design in three watersheds in the redwood forests of northern California, USA. Thinning treatments were intended to reduce canopy closure or basal area within the riparian zone by up to 50% on both sides of the stream channel along a 100-200 m stream reach. We found that responses to thinning ranged widely depending on the intensity of thinning treatments. In the watersheds with more intensive treatments, thinning reduced shade, increased light, and altered stream thermal regimes in thinned and downstream reaches. Thinning shifted thermal regimes by increasing maximum temperatures, thermal variability, and the frequency and duration of elevated temperatures. These thermal responses occurred primarily during summer but also extended into spring and fall. Longitudinal profiles indicated that increases in temperature associated with thinning frequently persisted downstream, but downstream effects depended on the magnitude of upstream temperature increases. Model selection analyses indicated that local changes in shade as well as upstream thermal conditions and proximity to upstream treatments explained variation in stream temperature responses to thinning. In contrast, in the study watershed with less intensive thinning, smaller changes in shade and light resulted in minimal stream temperature responses. Collectively, our data shed new light on the stream thermal responses to riparian thinning. These results provide relevant information for managers considering thinning as a viable restoration strategy for second-growth riparian forests.

New 3D measurements of large redwood trees for biomass and structure.

Large trees are disproportionately important in terms of their above ground biomass (AGB) and carbon storage, as well as their wider impact on ecosystem structure. They are also very hard to measure and so tend to be underrepresented in measurements and models of AGB. We show the first detailed 3D terrestrial laser scanning (TLS) estimates of the volume and AGB of large coastal redwood Sequoia sempervirens trees from three sites in Northern California, representing some of the highest biomass ecosystems on Earth. Our TLS estimates agree to within 2% AGB with a species-specific model based on detailed manual crown mapping of 3D tree structure. However TLS-derived AGB was more than 30% higher compared to widely-used general (non species-specific) allometries. We derive an allometry from TLS that spans a much greater range of tree size than previous models and so is potentially better-suited for use with new Earth Observation data for these exceptionally high biomass areas. We suggest that where possible, TLS and crown mapping should be used to provide complementary, independent 3D structure measurements of these very large trees.

Eliciting callus culture for production of hepatoprotective flavonoids and phenolics from Sequoia sempervirens (D. Don Endl).

The aim of our study is to estimate the hepatoprotective effects of the ethanolic extract of the leaves of Sequoia sempervirens by determination of liver biomarkers (ALT, AST, total bilirubin and albumin in serum) and by histopathological examinations using thioacetamide-induced (TAA) liver injury model. Concurrent administration of ethanolic extracts of S. sempervirens leaves improved the alterations in liver morphology where it was a potent protector of the liver. The potential of L-phenylalanine and silver nitrate as chemical elicitors as well as UV radiation as a physical elicitor on flavonoid production in callus culture of S. sempervirens were emphasized. Murashige and Skoog's medium fortified with phenylalanine and silver nitrate enhanced the production of flavonoids and phenolics. HPLC analysis was performed for qualitative and quantitative estimation of some flavonoid compounds in the produced calli in comparison with the mother plant. This finding highlights the potential use of S. sempervirens in the treatment of liver dysfunction.

The effect of low- and high-power microwave irradiation on in vitro grown Sequoia plants and their recovery after cryostorage.

Two distinct microwave power levels and techniques have been studied in two cases: low-power microwave (LPM) irradiation on in vitro Sequoia plants and high-power microwave (HPM) exposure on recovery rates of cryostored (-196°C) Sequoia shoot apices. Experimental variants for LPM exposure included: (a) in vitro plants grown in regular conditions (at 24 ± 1°C during a 16-h light photoperiod with a light intensity of 39.06 µEm-2 s-1 photosynthetically active radiation), (b) in vitro plants grown in the anechoic chamber with controlled environment without microwave irradiation, and (c) in vitro plants grown in the anechoic chamber with LPM irradiation for various times (5, 15, 30, 40 days). In comparison to control plants, significant differences in shoot multiplication and growth parameters (length of shoots and roots) were observed after 40 days of LPM exposure. An opposite effect was achieved regarding the content of total soluble proteins, which decreased with increasing exposure time to LPM. HPM irradiation was tested as a novel rewarming method following storage in liquid nitrogen. To our knowledge, this is the first report using this type of rewarming method. Although, shoot tips subjected to HPM exposure showed 28% recovery following cryostorage compared to 44% for shoot tips rewarmed in liquid medium at 22 ± 1 °C, we consider that the method represent a basis and can be further improved. The results lead to the overall conclusion that LPM had a stimulating effect on growth and multiplication of in vitro Sequoia plants, while the HPM used for rewarming of cryopreserved apices was not effective to achieve high rates of regrowth after liquid nitrogen exposure.

Contrasting drought-response strategies in California redwoods.

We compared the physiology and growth of seedlings originating from different Sequoia sempervirens (D. Don.) Endl. (coast redwood) and Sequoiadendron giganteum (Lindl.) Buchh. (giant sequoia) populations subjected to progressive drought followed by a recovery period in a controlled greenhouse experiment. Our objective was to examine how multiple plant traits interact to influence the response of seedlings of each species and seed population to a single drought and recovery cycle. We measured soil and plant water status, leaf gas exchange, stem embolism and growth of control (well-watered) and drought-stressed (water withheld) seedlings from each population at the beginning, middle and end of a 6-week drought period and again 2 weeks after re-watering. The drought had a significant effect on many aspects of seedling performance, but water-stressed seedlings regained most physiological functioning by the end of the recovery period. Sequoiadendron seedlings exhibited a greater degree of isohydry (water status regulation), lower levels of stem embolism, higher biomass allocation to roots and lower sensitivity of growth to drought compared with Sequoia. Only minor intra-specific differences were observed among populations. Our results show that seedlings of the two redwood species exhibit contrasting drought-response strategies that align with the environmental conditions these trees experience in their native habitats, and demonstrate trade-offs and coordination among traits affecting plant water use, carbon gain and growth under drought.

Millennium-scale crossdating and inter-annual climate sensitivities of standing California redwoods.

Extremely decay-resistant wood and fire-resistant bark allow California's redwoods to accumulate millennia of annual growth rings that can be useful in biological research. Whereas tree rings of Sequoiadendron giganteum (SEGI) helped formalize the study of dendrochronology and the principle of crossdating, those of Sequoia sempervirens (SESE) have proven much more difficult to decipher, greatly limiting dendroclimatic and other investigations of this species. We overcame these problems by climbing standing trees and coring trunks at multiple heights in 14 old-growth forest locations across California. Overall, we sampled 1,466 series with 483,712 annual rings from 120 trees and were able to crossdate 83% of SESE compared to 99% of SEGI rings. Standard and residual tree-ring chronologies spanning up to 1,685 years for SESE and 1,538 years for SEGI were created for each location to evaluate crossdating and to examine correlations between annual growth and climate. We used monthly values of temperature, precipitation, and drought severity as well as summer cloudiness to quantify potential drivers of inter-annual growth variation over century-long time series at each location. SESE chronologies exhibited a latitudinal gradient of climate sensitivities, contrasting cooler northern rainforests and warmer, drier southern forests. Radial growth increased with decreasing summer cloudiness in northern rainforests and a central SESE location. The strongest dendroclimatic relationship occurred in our southernmost SESE location, where radial growth correlated negatively with dry summer conditions and exhibited responses to historic fires. SEGI chronologies showed negative correlations with June temperature and positive correlations with previous October precipitation. More work is needed to understand quantitative relationships between SEGI radial growth and moisture availability, particularly snowmelt. Tree-ring chronologies developed here for both redwood species have numerous scientific applications, including determination of tree ages, accurate dating of fire-return intervals, archaeology, analyses of stable isotopes, long-term climate reconstructions, and quantifying rates of carbon sequestration.

Structural development of redwood branches and its effects on wood growth.

Redwood branches provide all the carbohydrates for the most carbon-heavy forests on Earth, and recent whole-tree measurements have quantified trunk growth rates associated with complete branch inventories. Providing all of a tree's photosynthetic capacity, branches represent an increasing proportion of total aboveground wood production as trees enlarge. To examine branch development and its effects on wood volume growth, we dissected 31 branches from eight Sequoia sempervirens (D. Don) Endl. and seven Sequoiadendron giganteum Lindl. trees. The cambium-area-to-leaf-area ratio was maintained with size and age but increased with light availability, whereas the heartwood-deposition-area-to-leaf-area ratio increased with size and age but was insensitive to light availability. The proportion of foliage mass arrayed in <1-cm-diameter epicormic shoots increased with decreasing light and was higher in Sequoia (20-60%) than in Sequoiadendron (3-16%). Well-illuminated branches concentrated leaves higher and distally, while shaded branches distributed leaves lower and proximally. In similar light environments, older branches distributed leaves lower and more proximally than younger branches. Branch size, light, species, heartwood area, a heartwood-area-species interaction, and ovulate cone mass predicted 87.5% of the variability in wood volume growth of branches. After accounting for the positive effects of size and light, wood volume growth declined with heartwood area and age. The effect of age was trivial compared to the effect of heartwood area, suggesting that heartwood expansion caused the age-related decline in wood volume growth. Additionally, Sequoiadendron branches of similar size and light environment with more ovulate cones produced less wood, even though these cones were long-lived and photosynthetic, reflecting the energetic cost of seed production. These results contributed to a conceptual model of branch development in which light availability, injury, heartwood content, gravity, and time interact to produce the high degree of branch structural variation evident within redwood crowns.

The right tree for the job? perceptions of species suitability for the provision of ecosystem services.

Stakeholders in plantation forestry are increasingly aware of the importance of the ecosystem services and non-market values associated with forests. In New Zealand, there is significant interest in establishing species other than Pinus radiata D. Don (the dominant plantation species) in the belief that alternative species are better suited to deliver these services. Significant risk is associated with this position as there is little objective data to support these views. To identify which species were likely to be planted to deliver ecosystem services, a survey was distributed to examine stakeholder perceptions. Stakeholders were asked which of 15 tree attributes contributed to the provision of five ecosystem services (amenity value, bioenergy production, carbon capture, the diversity of native habitat, and erosion control/water quality) and to identify which of 22 candidate tree species possessed those attributes. These data were combined to identify the species perceived most suitable for the delivery of each ecosystem service. Sequoia sempervirens (D.Don) Endl. closely matched the stakeholder derived ideotypes associated with all five ecosystem services. Comparisons to data from growth, physiological and ecological studies demonstrated that many of the opinions held by stakeholders were inaccurate, leading to erroneous assumptions regarding the suitability of most candidate species. Stakeholder perceptions substantially influence tree species selection, and plantations established on the basis of inaccurate opinions are unlikely to deliver the desired outcomes. Attitudinal surveys associated with engagement campaigns are essential to improve stakeholder knowledge, advancing the development of fit-for-purpose forest management that provides the required ecosystem services.

Increased susceptibility to drought-induced mortality in Sequoia sempervirens (Cupressaceae) trees under Cenozoic atmospheric carbon dioxide starvation.

PREMISE OF THE STUDY: Climate-induced forest retreat has profound ecological and biogeochemical impacts, but the physiological mechanisms underlying past tree mortality are poorly understood, limiting prediction of vegetation shifts with climate variation. Climate, drought, fire, and grazing represent agents of tree mortality during the late Cenozoic, but the interaction between drought and declining atmospheric carbon dioxide ([CO2]a) from high to near-starvation levels ∼34 million years (Ma) ago has been overlooked. Here, this interaction frames our investigation of sapling mortality through the interdependence of hydraulic function, carbon limitation, and defense metabolism. • METHODS: We recreated a changing Cenozoic [CO2]a regime by growing Sequoia sempervirens trees within climate-controlled growth chambers at 1500, 500, or 200 ppm [CO2]a, capturing the decline toward minimum concentrations from 34 Ma. After 7 months, we imposed drought conditions and measured key physiological components linking carbon utilization, hydraulics, and defense metabolism as hypothesized interdependent mechanisms of tree mortality. • KEY RESULTS: Catastrophic failure of hydraulic conductivity, carbohydrate starvation, and tree death occurred at 200 ppm, but not 500 or 1500 ppm [CO2]a. Furthermore, declining [CO2]a reduced investment in carbon-rich foliar defense compounds that would diminish resistance to biotic attack, likely exacerbating mortality. • CONCLUSIONS: Low-[CO2]a-driven tree mortality under drought is consistent with Pleistocene pollen records charting repeated Californian Sequoia forest contraction during glacial periods (180-200 ppm [CO2]a) and may even have contributed to forest retreat as grasslands expanded on multiple continents under low [CO2]a over the past 10 Ma. In this way, geologic intervals of low [CO2]a coupled with drought could impose a demographic bottleneck in tree recruitment, driving vegetation shifts through forest mortality.

Small RNAs of Sequoia sempervirens during rejuvenation and phase change.

In this work, the population of small RNAs (sRNAs) was studied in the gymnosperm Sequoia sempervirens during phase changes, specifically in the juvenile, adult and rejuvenated plants obtained in vitro. The potential target genes of Sequoia sRNAs were predicted through bioinformatics. Rejuvenation is a pivotal process in woody plants that enables them to regain their growth potential, which results in the recovery of physiologic and molecular characteristics that were lost when the juveniles mature into adult plants. The results from the five repeated graftings of juvenile, adult and rejuvenated plants in vitro showed that sRNAs could be classified into structural RNAs (Group I), small interfering RNAs (Group II), annotated microRNAs (Group III, and unannotated sRNAs (Group IV). The results indicate that only 573 among 15,485,415 sRNAs (Groups III and IV) had significantly different expression patterns associated with rejuvenation and phase change. A total of 215 sRNAs exhibited up-regulated expression patterns in adult shoots, and 358 sRNAs were down-regulated. Expression profiling and prediction of possible target genes of these unique small RNAs indicate possible functions in the control of photosynthetic efficiency and rooting competence abundance during plant rejuvenation. Moreover, the increase in SsmiR156 and decrease in SsmiR172 during plant rejuvenation suggested that these two microRNAs extensively affect phase transition.

Designing the ideal habitat for entomopathogen use in nursery production.

BACKGROUND: Greenhouse and nursery producers use entomopathogens (nematodes and fungi) to control soil pests. Although it is known that the physical and chemical properties of mineral soil significantly impact upon soil pathogens, the influence of soilless media used for plant production on entomopathogen performance is poorly understood. RESULTS: Survival and foraging distance were differently affected by sand:peat, bark and sawdust media for entomopathogenic nematodes, but not for the immobile fungus Metarhizium anisopliae. Redwood sawdust medium consistently had a negative impact upon entomopathogenic nematodes. Dividing media into individual components supported the hypothesis that redwood sawdust reduced foraging and infection abilities of S. riobrave and H. bacteriophora. Physically altering the components by adding sand significantly improved foraging and infection success for S. riobrave in media not optimum for foraging. CONCLUSION: This study is the first to highlight the importance of selecting the appropriate soilless media and pathogen species combinations to increase efficacy of biological control. H. bacteriophora was able to find hosts in a wider diversity of medium components than S. riobrave, although both nematode species performed well in peat moss and recycled plant material. These results suggest that peat moss, recycled plant material and hardwood bark are components amenable to EPN biological control programs.

Cryopreservation of redwood (Sequoia sempervirens) in vitro buds using vitrification-based techniques.

In this study, the efficiency of three vitrification-based cryopreservation techniques, i.e. vitrification, encapsulation-vitrification and droplet-vitrification were compared for cryopreserving Sequoia sempervirens apical and basal buds sampled from in vitro shoot cultures. The effect of cold-hardening of mother-plants and of bud culture medium and sucrose preculture was also investigated. Culture of apical and basal buds sampled from cold-hardened mother-plants on Quoirin and Lepoivre medium with activated charcoal had a positive effect on regrowth. Only droplet-vitrification ensured survival and regrowth after cryopreservation. After cryopreservation, regeneration of apical buds was possible for PVS2 exposure durations between 90 and 180 min but it remained low, with a maximum of 18 percent after 135 min treatment. With basal buds, regeneration after cryopreservation was possible over a larger range of PVS2 treatment durations, between 30 and 180 min. The highest regeneration percentage was slightly higher (22 percent) than that measured with apical buds, and was also achieved after 135 min PVS2 exposure.

Effects of height on treetop transpiration and stomatal conductance in coast redwood (Sequoia sempervirens).

Treetops become increasingly constrained by gravity-induced water stress as they approach maximum height. Here we examine the effects of height on seasonal and diurnal sap flow dynamics at the tops of 12 unsuppressed Sequoia sempervirens (D. Don) Endl. (coast redwood) trees 68-113 m tall during one growing season. Average treetop sap velocity (V(S)), transpiration per unit leaf area (E(L)) and stomatal conductance per unit leaf area (G(S)) significantly decreased with increasing height. These differences in sap flow were associated with an unexpected decrease in treetop sapwood area-to-leaf area ratios (A(S):A(L)) in the tallest trees. Both E(L) and G(S) declined as soil moisture decreased and vapor pressure deficit (D) increased throughout the growing season with a greater decline in shorter trees. Under high soil moisture and light conditions, reference G(S) (G(Sref); G(S) at D = 1 kPa) and sensitivity of G(S) to D (-δ; dG(S)/dlnD) significantly decreased with increasing height. The close relationship we observed between G(Sref) and -δ is consistent with the role of stomata in regulating E(L) and leaf water potential (Ψ(L)). Our results confirm that increasing tree height reduces gas exchange of treetop foliage and thereby contributes to lower carbon assimilation and height growth rates as S. sempervirens approaches maximum height.

Climatic context and ecological implications of summer fog decline in the coast redwood region.

Biogeographical, physiological, and paleoecological evidence suggests that the coast redwood [Sequoia sempervirens (D. Don) Endl.] is closely associated with the presence of summer marine fog along the Pacific coast of California. Here we present a novel record of summer fog frequency in the coast redwood region upon the basis of direct hourly measurements of cloud ceiling heights from 1951 to 2008. Our analysis shows that coastal summer fog frequency is a remarkably integrative measure of United States Pacific coastal climate, with strong statistical connections to the wind-driven upwelling system of the California Current and the broad ocean temperature pattern known as the Pacific Decadal Oscillation. By using a long-term index of daily maximum land temperatures, we infer a 33% reduction in fog frequency since the early 20th century. We present tree physiological data suggesting that coast redwood and other ecosystems along the United States west coast may be increasingly drought stressed under a summer climate of reduced fog frequency and greater evaporative demand.

Hydrostatic constraints on morphological exploitation of light in tall Sequoia sempervirens trees.

We studied changes in morphological and physiological characteristics of leaves and shoots along a height gradient in Sequoia sempervirens, the tallest tree species on Earth, to investigate whether morphological and physiological acclimation to the vertical light gradient was constrained by hydrostatic limitation in the upper crown. Bulk leaf water potential (Psi) decreased linearly and light availability increased exponentially with increasing height in the crown. During the wet season, Psi was lower in the outer than inner crown. C isotope composition of leaves (delta(13)C) increased with increasing height indicating greater photosynthetic water use efficiency in the upper crown. Leaf and shoot morphology changed continuously with height. In contrast, their relationships with light availability were discontinuous: morphological characteristics did not correspond to increasing light availability above 55-85 m. Mass-based chlorophyll concentration (chl) decreased with increasing height and increasing light availability. In contrast, area-based chl remained constant or increased with increasing height. Mass-based maximum rate of net photosynthesis (P (max)) decreased with increasing height, whereas area-based P (max) reached maximum at 78.4 m and decreased with increasing height thereafter. Mass-based P (max) increased with increasing shoot mass per area (SMA), whereas area-based P (max) was not correlated with SMA in the upper crown. Our results suggest that hydrostatic limitation of morphological development constrains exploitation of light in the upper crown and contributes to reduced photosynthetic rates and, ultimately, reduced height growth at the tops of tall S. sempervirens trees.

A hydraulic-photosynthetic model based on extended HLH and its application to Coast redwood (Sequoia sempervirens).

Although recent investigations [Ryan, M.G., Yoder, B.J., 1997. Hydraulic limits to tree height and tree growth. Bioscience 47, 235-242; Koch, G.W., Sillett, S.C.,Jennings, G.M.,Davis, S.D., 2004. The limits to tree height. Nature 428, 851-854; Niklas, K.J., Spatz, H., 2004. Growth and hydraulic (not mechanical) constraints govern the scaling of tree height and mass. Proc. Natl Acad. Sci. 101, 15661-15663; Ryan, M.G., Phillips, N., Bond, B.J., 2006. Hydraulic limitation hypothesis revisited. Plant Cell Environ. 29, 367-381; Niklas, K.J., 2007. Maximum plant height and the biophysical factors that limit it. Tree Physiol. 27, 433-440; Burgess, S.S.O., Dawson, T.E., 2007. Predicting the limits to tree height using statistical regressions of leaf traits. New Phytol. 174, 626-636] suggested that the hydraulic limitation hypothesis (HLH) is the most plausible theory to explain the biophysical limits to maximum tree height and the decline in tree growth rate with age, the analysis is largely qualitative or based on statistical regression. Here we present an integrated biophysical model based on the principle that trees develop physiological compensations (e.g. the declined leaf water potential and the tapering of conduits with heights [West, G.B., Brown, J.H., Enquist, B.J., 1999. A general model for the structure and allometry of plant vascular systems. Nature 400, 664-667]) to resist the increasing water stress with height, the classical HLH and the biochemical limitations on photosynthesis [von Caemmerer, S., 2000. Biochemical Models of Leaf Photosynthesis. CSIRO Publishing, Australia]. The model has been applied to the tallest trees in the world (viz. Coast redwood (Sequoia sempervirens)). Xylem water potential, leaf carbon isotope composition, leaf mass to area ratio at different heights derived from the model show good agreements with the experimental measurements of Koch et al. [2004. The limits to tree height. Nature 428, 851-854]. The model also well explains the universal trend of declining growth rate with age.

Shoot regeneration and somatic embryogenesis from needles of redwood (Sequoia sempervirens (D.Don.) Endl.).

A rapid and effective system of somatic embryogenesis and organogenesis from the in vitro needles of redwood (Sequoia sempervirens (D.Don.) Endl.) had been established. The influences of plant growth regulators (PGRs) and days of seedlings in vitro on adventitious bud regeneration and somatic embryogenesis were studied. The process of somatic embryo formation was also observed. The results showed that embryogenic callus was induced and proliferated on Schenk and Hildebrandt (SH) medium with BA (0.5 mg/l), KT (0.5 mg/l) and IBA (1.0 mg/l). SH medium containing BA (0.5 mg/l), KT (0.2 mg/l) and IBA (0.2 mg/l) effectively promoted adventitious bud regeneration. The highest frequency (66.3%) of direct somatic embryogenesis was obtained in the combination of BA (0.5 mg/l) and IBA (0.5 mg/l). The optimal days of seedling in vitro for adventitious bud and somatic embryogenesis were 30 days and 30-40 days, respectively. The developments of somatic embryos were similar to that of zygotic embryogenesis. The result of histocytological studies indicated that proteins were gradually accumulated in the process of somatic embryo formation and there were two peaks of starch grains accumulation that one was in the embryogenic callus and the other was in the globular embryos. These results indicated that starch and protein were closely related with the energy supply and the molecular base of somatic embryogenesis, respectively.

Carbon content variation in boles of mature sugar maple and giant sequoia.

At present, a carbon (C) content of 50% (w/w) in dry wood is widely accepted as a generic value; however, few wood C measurements have been reported. We used elemental analysis to investigate C content per unit of dry matter and observed that it varied both radially and vertically in boles of two old-growth tree species: sugar maple (Acer saccharum Marsh.) and giant sequoia (Sequoiadendron giganteum (Lindl.) Bucholz). In sugar maple there was considerable variation in tree ring widths among four radii for particular annual layers of xylem, revealing that the annual rate of C assimilation differs around the circumference and from the base of each tree to its top, but the observed variation in C content was unrelated to diameter growth rate and strongly related to the calendar year when the wood was formed. Carbon content in sugar maple wood increased in an approximately linear fashion, from < 50 to 51% from pith to cambium, at both the base and top of the boles. In giant sequoia, C was essentially constant at > 55% across many hundreds of years of heartwood, but it declined abruptly at the sapwood-heartwood boundary and remained lower in all sapwood samples, an indication that heartwood formation involves anabolic metabolism. Factors that may be responsible for the different C contents and trends with age between sugar maple and sequoia trees are considered. Tree-ring data from this study do not support some of the key assumptions made by dendrochronology.

The limits to tree height.

Trees grow tall where resources are abundant, stresses are minor, and competition for light places a premium on height growth. The height to which trees can grow and the biophysical determinants of maximum height are poorly understood. Some models predict heights of up to 120 m in the absence of mechanical damage, but there are historical accounts of taller trees. Current hypotheses of height limitation focus on increasing water transport constraints in taller trees and the resulting reductions in leaf photosynthesis. We studied redwoods (Sequoia sempervirens), including the tallest known tree on Earth (112.7 m), in wet temperate forests of northern California. Our regression analyses of height gradients in leaf functional characteristics estimate a maximum tree height of 122-130 m barring mechanical damage, similar to the tallest recorded trees of the past. As trees grow taller, increasing leaf water stress due to gravity and path length resistance may ultimately limit leaf expansion and photosynthesis for further height growth, even with ample soil moisture.