DOI: 10.5061/dryad.k6djh9w3x
title: Data from: Decoupling between growth rate and storage remobilization in broadleaf temperate tree species

description: Temperate trees rely on carbon (C) and nutrient remobilisation from
 storage to resume growth after winter. Minimum storage levels during the
 growing season suggest that remobilisation could signify that C
 availability is insufficient to meet growth demands; consequently, growth
 might be C and/or nutrient limited. However, it remains unclear whether
 higher growth demands are covered by higher remobilization. This study
 examined whether higher C and nutrient demands associated with fast growth
 or deciduousness, rely on greater remobilisation. In 11 sympatric
 deciduous and evergreen angiosperm tree species from southern South
 América, the magnitude of seasonal remobilisation of C and nutrient
 storage was assessed as the seasonal minimums (relative to seasonal
 maximums) of whole tree non-structural carbohydrates (NSC), nitrogen (N),
 and phosphorus (P) concentrations and pools. The basal area increment and
 stem wood density were determined for each tree, from which the biomass
 radial increment (BRI) was estimated. The effects of leaf habit and BRI on
 the seasonal minimums NSCs and nutrient concentrations and pools were
 analysed using linear mixed-effects models. Radial growth was not related
 to seasonal minimum NSC or nutrient concentrations and pools in either the
 evergreens or deciduous angiosperms; thus, faster growth was not
 associated with greater remobilisation of C or nutrient stores. Further,
 larger trees grew faster than smaller ones, but did not have higher
 remobilisation. Deciduous species had higher year-round whole tree NSC and
 nutrient concentrations than evergreens; however, both groups had similar
 BRI and seasonal minimum concentrations and pools of NSCs and nutrients.
 Neither growth rate nor leaf habit drove the magnitude of C and nutrient
 remobilisation in the angiosperm trees examined here, indicating no C or
 nutrient limitation. This result contradicts the view that growth and
 storage strongly regulate one another, as proposed by a growth-storage
 trade-off.
description: Seasonal sampling of the dominant tree species of four
 cold-temperate forests was performed in Patagonia, specifically in the
 Aysén region of Chile. The sites were: Aiken (45°27´S, 72°44´W, 45 m
 a.s.l.), Coyhaique (45°32´S, 72°03´W, 466 m a.s.l.), Tranquilo (K10)
 (46°38´S, 72°47´W, 331 m a.s.l.) and Exploradores (46°29´S, 73°09´W, 137 m
 a.s.l.). In each forest, six unshaded and healthy
 juvenile trees of two winter deciduous species (<em>Nothofagus
 antarctica</em> and <em>Ribes magellanicum, Na and Rm,
 respectively</em>) and two to five evergreen angiosperm species
 (<em>Nothofagus nitida (Nn), Embothrium coccineum (Ec), Aristotelia
 chilensis (Ac), Myrceugenia planipes (Mp), Luma apiculata (La), Nothofagus
 dombeyi (Nd), Rhaphithamnus spinosus (Ra), Nothofagus betuloides (Nb),
 Drimys winteri (Dw</em>)) were selected for chemical analyses and to
 measure radial growth. Coarse roots, stems, branches, twigs, and new
 leaves were sampled between 10:00 and 16:00 h at a single time point
 (hereafter referred to as “sampling date”), which corresponded with
 mid-spring, mid-summer, mid-autumn, and mid-winter. For each sampling
 date, the four forests were sampled over a 3-week period.
 One to-the-pith long stem core at breast height (1.35 m) was
 sampled using a 5.15 mm increment borer for NSC analyses. Another stem
 core was similarly sampled on the first sampling date to determine tree
 growth. Growth was measured as the increment in basal area of the last 6
 years (BAI<sub>6</sub>). To account for potential differences
 in biomass investment due to wood density, BAI<sub>6</sub> was
 converted into the biomass radial increment (BRI), which represents the
 biomass increment of a hollow cylinder of 1 cm height, for which the wall
 thickness (i.e. the difference between outer and inner diameter) is the
 BAI<sub>6</sub>. The volume of this cylinder was then
 multiplied by the species wood density of a given species to obtain the
 biomass of the cylinder. All samples for chemical
 analyses were placed in a forced-air stove at 65 °C for 72 h. The material
 was then ground into a fine powder and stored under cool, dry, and dark
 conditions, until chemical analyses were conducted.
 Carbohydrate concentrations were analysed for all the samples. C
 reserves were assessed by determining NSC; specifically, as the sum of the
 three most abundant low molecular-weight soluble sugars (glucose, fructose
 and sucrose; <em>hereafter</em> SGF) and starch. NSC
 concentrations were analysed following the procedure of Hoch et al. (2002)
 with some modifications as described by Piper et al. (2019). Nitrogen (N)
 concentrations were determined from 0.2 g of dried ground material using a
 combustion analyser (LECO TruSpec Micro CHN, Centro de Investigación de
 Ecosistemas de la Patagonia, Coyhaique, Chile). Inorganic total phosphate
 (P) concentrations were determined from 100 mg dried ground material based
 on the procedure described by Murphy & Riley (1962).
 The dry wood biomass of each organ was estimated using previously
 developed allometric functions. For each sampled tree and on each date,
 the NSC, starch, SGF, N, and P pools were first calculated per each organ
 as the product between the biomass and the concentration for each organ.
 Then, the pools of the different organs of a tree were summed to estimate
 the whole-tree total pool on each sampling date. The seasonal minimum pool
 of NSC, starch, SGF, N, and P was determined for each tree on the four
 sampling dates (as the lowest value), and was expressed relative to the
 seasonal maximum pool (percentage). Thus, the higher the seasonal minimum,
 the lower the amplitude of seasonal variation in a given pool (Fig. 1).
 Similar calculations were performed for NSC, starch, SGF, N, and P
 concentrations; however, in these cases, seasonal minimums were estimated
 separately for each organ.
description: There are some missing values of N and P
 concentrations. Wood density was measured for most
 sites in this study, although for a few cases (species in a given site) I
 used published data available elsewhere. 
description: # Data from: Decoupling between growth rate and storage remobilization in
 broadleaf temperate tree species
 Seasonal sampling of
 leaves, stem sapwood, twigs, branch sapwood and root sapwood of the
 dominant tree species of four cold temperate forests was performed in
 Patagonia, Aysén region of Chile, for determination of concentrations of
 non-structural carbohydrates (NSC, in %), starch (St, in %), SGF (sucrose,
 glucose and fructose, in %), nitrogen (N, in %), phosphorus (P, in
 mg.g-1). Also, wood density (WD), and specific leaf area (SLA) was
 measured. Allometric equations taken from literature were used to estimate
 whole-tree pools of NSC and nutrients. ## Description of the data and file
 structure **​File “Carbohydrates and nutrients concentrations and pools“**
 1. Sampling sites. Aiken: Parque Aiken del Sur, 45°27´S, 72°44´W, 45 m
 a.s.l. RNC: Reserva Nacional Coyhaique, 45°32´S, 72°03´W, 466 m a.s.l.
 K10: Puerto Tranquilo, 46°38´S, 72°47´W, 331 m a.s.l. Exploradores: Bahía
 exploradores, 46°29´S, 73°09´W, 137 m a.s.l.Sampling month 2. Tree Organ
 selected for sampling 3. Species (11 species sampled, 3 of them common for
 the four sites). Ac: Aristotelia chilensis Nn: Nothofagus nitida Na: N.
 antarctica Nb: N. betuloides Nd: N. dombeyi La: Luma apiculata Mp:
 Myrceugenia planipes Dw: Drimys winteri Rs: Rhaphithamnus spinosus  Ec:
 Embothrium coccineum Rm: Ribes magellanicum 4. Individual (6 individuals
 were sampled per site, month and species). 5. NSC (%): Concentration of
 non-structural carbohydrates, in % of dry weight 6. St: concentration of
 starch (in %) 7. SGF: Concentration of glucose, fructose and glucose (in
 %) 8. N (%): Concentration of nitrogen (as % of dry weight) 9. P:
 Concentration of phosphorus, in mg per gram of dry weight 10. Organ
 weight: the weight of each tree organ estimated from allometric functions
 presented in sheet “allometric functions“ of the same file, in Kg 11. NSC
 pool: Content of non-structural carbohydrates, calculated as organ NSC
 concentration x organ biomass 12. St pool: Content of starch, calculated
 as organ starch concentration x organ biomass 13. SGF pool: Content of
 sucrose, glucose and fructose, calculated as organ SGF concentration x
 organ biomass 14. N pool: Nitrogen content, calculated as N organ
 concentration x organ biomass 15. P pool: Phosphorus content, calculated
 as P organ concentration x organ biomass **​File “SLA and WD seasonal“**
 1. Site: Study Site. Aiken: Parque Aiken del Sur, 45°27´S, 72°44´W, 45 m
 a.s.l. RNC: Reserva Nacional Coyhaique, 45°32´S, 72°03´W, 466 m a.s.l.
 K10: Puerto Tranquilo, 46°38´S, 72°47´W, 331 m a.s.l. Exploradores: Bahía
 exploradores, 46°29´S, 73°09´W, 137 m a.s.l. 2. Species: Study Species, as
 defined in the metadata sheet. Ac: Aristotelia chilensis Nn: Nothofagus
 nitida Na: N. antarctica Nb: N. betuloides Nd: N. dombeyi La: Luma
 apiculata Mp: Myrceugenia planipes Dw: Drimys winteri Rs: Rhaphithamnus
 spinosus  Ec: Embothrium coccineum Rm: Ribes magellanicum 3. Individual:
 Tree ID. From 1 to 6 4. Leaf weight  (in grams): Dry mass of a group of
 leaves photographed on a white sheet, for specific leaf mass calculation.
 Values are given for November, February, and July 5.  Leaf Area (in cm2):
 Leaf area of group of leaves photographed on a white sheet, for specific
 leaf mass calculation. Values are given for November, February, and July
 6. SLA: Specific leaf area: Leaf area (cm2) by leaf weight (g)