1. Introduction
Changes in environmental conditions due to climate change influence ecological systems. Particularly the projected temperature rise for the next decades may result in a large variation of possible impacts on forest ecosystems [
1]. Various model studies have examined the response of forest growth to changing climate conditions to develop future forest management strategies ensuring ecologically viable forest ecosystems [
2,
3,
4,
5].
Resistance to drought stress is an important factor for the functioning of many forest ecosystems, because of the close relationship between carbon and water cycles in trees expressed by the linkage of stomatal conductance and photosynthesis [
6,
7]. Climate warming might lead to decreasing water availability due to higher evapotranspiration rates in most European regions [
8]. In Mediterranean forests, the morphology, phenology and physiology of trees are well-adapted to recurring drought events [
9]. In Central Europe, severe droughts are less frequent and thus the adaptation to drought stress is less distinctive. Especially, the growth of mature forest stands is strongly dependent on water availability during the growing period [
10].
In the federal state of Brandenburg, Germany, 70% of forest area is stocked with pure Scots pine (
Pinus sylvestris L.) stands which are partly being converted to mixed Scots pine-oak stands (
Quercus petraea [Matt.] Liebl.). The region is characterized by dry climate conditions and soils with low water storage capacity. The vulnerability of Scots pine to severe drought events has already been subject to discussion [
11,
12]. In addition, the stability of pure Scots pine stands is unclear at the southern and western borders of the distribution area under the condition of continuous temperature increase in the next decades [
13].
Therefore, assessing the magnitude of severe drought impacts in the future is essential. Doing so requires the realistic mapping of plant available water, which represents the water supply to tree roots. There are two theories on water flow within trees. First, the
cohesion-tension theory of the ascent of sap deals with the physics of the sap water movement [
14]. Second, the
electrical analogy describes the water transport within the soil-plant-water continuum by using resistances, capacitances, and water potentials [
14].
Process-based forest growth models are suitable scientific tools to study climate impacts and tree responses to droughts [
15,
16,
17,
18], although drought stress and especially its implications for tree mortality are often not sufficiently covered in these models [
19]. Forest canopy models for the soil-plant-atmosphere-continuum as developed by Williams
et al. [
20] link carbon uptake of forest stands to weather conditions and soil properties. They can provide reliable information about the relationships between carbon dioxide uptake, stomatal conductance and transpiration. The water flow is described as an electrical circuit and water uptake of roots is simulated by calculating water flow dependent on the water potential gradient from soil to atmosphere [
21,
22,
23,
24,
25,
26]. The considered water potentials and resistances depend nonlinearly on the water contents in the particular components along the soil-plant-atmosphere continuum (SPAC). The maximum rate of water supply by roots is determined by the minimum sustainable leaf water potential, the soil conductivity for water flow and the root resistance to water uptake. This approach permits a species-specific characterization of belowground competition in mixed forest stands since modeling of resistances of root water uptake directly depends on relative fine root length densities, a species-specific, measureable parameter. The SPAC-model predicts varying stomatal conductance so that daily carbon uptake is maximized within the limitations of canopy water and nitrogen availability.
However, the described model by Williams
et al. [
20] is less useful to analyze impacts of extreme weather events (e.g., droughts) on diameter increment and subsequently timber yield of forest stands. For this purpose, models are required that incorporate a carbon allocation module and consider stand development with regard to management and mortality. Economic and ecological sustainability under continuous climate warming can be investigated with these kinds of forest growth models. However, in these models usually a simple function accounts for limited water uptake at decreasing soil water content. They simulate water uptake with an empirical reduction function, which is linearly dependent on soil water content [
27,
28].
Knowledge about the performance of different water uptake model approaches within a physiological-based forest growth model helps to detect potential severe drought events under future climate scenarios. Furthermore, this knowledge is of major importance for the development of suitable tools for planning adaption strategies as for instance forest conversion. The objective of this paper is to identify the effect of two commonly used water uptake approaches on the simulated amount of plant available water and to assess the resulting effects on tree growth in pure pine and mixed oak-pine forests. Both approaches are implemented in the process based forest growth model 4C (FORESEE—FORESt Ecosystems in a changing Environment) [
2] and applied in a simulation study. This simulation study strives to respond to the following questions: (1) Are there differences between both approaches regarding the water supply from the roots? (2) How well can drought years, observed by annual tree ring analyses, be reproduced by the different water uptake approaches? (3) Is there an approach that performs best regarding different validation data sets?
Due to the complexity of comparing these two approaches with regard to direct and indirect effects on different model output variables (e.g., soil water limitation → reduction of transpiration → reduction of diameter increment), we here employ three different data sets to discuss the two model approaches at several time scales relevant for tree growth. The data sets include (1) daily transpiration and (2) soil water content measurements over a short-term period, which we use to directly evaluate root water uptake; and (3) annual tree ring data, serving as a long-term validation archive for severe drought events, which constitute an indirect link to root water uptake impacts [
29]. We initialize the model for each forest stand with the soil description available for that stand. Then, we run the model with the two different water uptake approaches for each stand and only the model formulation with regard to water uptake is different.
2. Material and Methods
2.1. Forest Growth Model 4C
The physiologically-based forest growth model 4C has been used to simulate growth, water, and carbon budget of trees and soils under current and projected climate condition and to analyze the long-term growth behavior of forest stands. 4C was applied in different studies and validated across a wide range of forest sites [
2,
5,
30,
31]. Modeled processes are based on eco-physiological experiments, long-term studies of stand development, and physiological relationships [
32] at tree and stand level. Establishment, growth, and mortality are explicitly modeled on individual patches for which homogeneity is assumed. The model simulates tree species composition, forest structure, leaf area index as well as ecosystem carbon and water balances. The cohort structure of simulated tree growth enables a detailed analysis from stand to single-tree level, whereas a cohort represents trees of same species, age and dimensions. The length of the growing season is provided by a species-specific phenological model with prohibitors and inhibitors [
33]. The photosynthesis submodel is based on the photosynthesis model of Haxeltine and Prentice [
32]. The photosynthesis is calculated under the assumption of unlimited water and nutrient supply and then reduced based on actual nutrient and water limitations. Reductions of photosynthesis by water are represented through a drought reduction factor
Rdrind, calculated as the ratio between soil water supply and tree water demand, thus following the physiological logic of the model. The model variable
Rdrind is not a static parameter but dynamically calculated in the model as a result of the link between photosynthesis and water balance of the tree. We analyze it in this study to compare both water uptake approaches.
For simulating water and carbon balances in the forest soil, a multilayered soil model is implemented in 4C. Water balances are calculated using a bucket model approach [
34,
35]. The surplus water above field capacity percolates out of the last soil layer and thus represents deep soil water infiltration. The soil column is divided into different layers with optional thickness according to the horizons of the soil profile. Each layer, the humus layer and the deeper mineral layers, is considered homogeneous concerning its physical and chemical parameters. Water content and soil temperature of each soil layer are estimated as functions of the soil parameters, air temperature, and stand precipitation and control the decomposition and mineralization of organic matter. The water input to the soil is equivalent to the throughfall (precipitation minus interception). The interception is determined by the interception storage and the potential evapotranspiration [
36]. The interception storage is calculated from a leaf area index that is weighted by a species-specific storage capacity. The calculation of the soil temperature of each soil layer is described by Grote and Suckow [
27].
2.2. Modeling Water Uptake
2.2.1. Empirical Water Uptake Approach—WU1
The water uptake approach (WU1) assumes the water uptake of a tree cohort to be controlled by the transpiration demand, the relative amount of fine roots of a tree cohort in a specific soil layer relative to the total amount of fine roots in this soil layer, the available water above the wilting point,
WsWP, and a reduction function [
37]. The reduction function
fres is based on the assumption that optimal water uptake only occurs if water content,
Ws, is in range of ±10% of the field capacity,
WsFC. Hence, WU1 follows a demand-approach that looks stepwise from the upper to the lower soil layer for the required amount of water. The water demand of a tree cohort is calculated from the potential evapotranspiration and the canopy conductance with regard to photosynthesis. This demand is met by the supply of soil water, calculated from water content of the soil layers reduced by a water content dependent reduction factor (
Figure 1).
Figure 1.
Schematic overview of the simulated components for WU1 and WU2. With Wupt and Etree = sum of water uptake of a tree over nl number of soil layers, Ws = soil water content, Rr = root resistance, Rs = soil resistance, ψL = leaf water potential, ψr = root water potential, ψs = soil water potential.
Figure 1.
Schematic overview of the simulated components for WU1 and WU2. With Wupt and Etree = sum of water uptake of a tree over nl number of soil layers, Ws = soil water content, Rr = root resistance, Rs = soil resistance, ψL = leaf water potential, ψr = root water potential, ψs = soil water potential.
The water supply by roots
Wcupt [mm] per soil layer is calculated by WU1 as follows:
where
Ws is the soil water content [mm] of the respective soil layer. The reduction factor
fres that implies the reduced water uptake at water contents besides the optimum is simulated as follows:
where
Wssat is the saturated water content of soil layer,
WsFC is the soil water content at field capacity and
WsWP is the soil water content at wilting point.
2.2.2. Process-Based Water Uptake Approach—WU2
The second approach for water uptake (WU2) was developed by Campbell [
21] and describes the water uptake process along a soil-plant-atmosphere continuum (SPAC). The algorithm has been implemented in 4C to simulate water uptake based on the water potential gradient between leaf and soil analogous to electricity flow. The water flows from soil through xylem into the leaves and evaporates through the stomata into the atmosphere.
Figure 1 illustrates the simulated time dynamic components within the applied water uptake approach. Two major resistances in the system are calculated: the soil resistance (Equation (7)) and the root endodermis resistance (Equation (4)). Axial resistances within roots and xylem resistances within the stem are not considered in the model and the leaf resistance is constant over time and tree species. The potential gradient with the lowest water potential in the leaves and the highest water potential in the soil is the driver of the water flow in trees. The root resistance in a soil layer is inversely proportional to the relative length of fine roots in that layer. In the following we show the key equations from the original source of WU2 [
21]. The list of parameters of WU2 is provided in Table S1.
WU2 calculates the water supply of the roots
Wcupt for cohort c with water content of the soil
Ws and transpiration of one tree from cohort
c, which is multiplied by the number of trees within the cohort
Ntreec.
Etree is the transpiration flow within the soil-plant-atmosphere continuum of the single tree of cohort c.
The transpiration or water uptake of a tree
Etree over all soil layers, without considering xylem resistance, is calculated from the soil water potentials ψ
si, the xylem water potential ψ
xri and the soil and root resistances
Rsi and
Rri of all soil layers
nl.
The first major resistance
Rri (root resistance) is directly proportional to the total root resistance
Rrtot and inversely proportional to the fine root length density
Li [m·m
−3] in soil layer
i.
The second major resistance
Rsi (soil resistance) is calculated once per day for every soil layer
i by a simplified approach from the fine root length density
Li and the conductivity of root (
kr) and soil (
ks).
with:
where
rr is the fine root radius, Δ
zi the thickness of soil layer
i and
n a dimensionless parameter. We calculated water uptake with low, medium and high total root resistance (WU2-low, WU2-medium, and WU2-high; Table S1). The soil water potentials were calculated on the base of pedotransfer functions from the actual water content [
38].
2.3. Forest Stands and Site Conditions
The simulation experiments were conducted on four different forest sites in Germany. We choose two mono-species Scots pine (Pinus sylvestris L.) stands (P1, P2) and two mixed Scots pine-oak (Quercus petraea [Matt.] Liebl.) stands (M1, M2) located in the lowlands of northeastern Germany. P1 and P2 are part of the network of long-term forest monitoring sites within the ICP Forest program of the UN-ECE.
For the short-term validation, we accessed initialization data for the P1 and P2 stand from 1994 (
Table 1). The simulation period for long-term validation starts in 1951. Since there were no observed stand data for the start year of simulation in 1951, we used available data of inventories from 2006 (M1, M2) and 2004 (P1, P2) to generate stands for 1951 from yield table data. We assumed stability in the site quality index (mean dominant tree height at age 100) and in the share of basal area in mixed stands and initialized the stands on the base of tree age, mean height, mean diameter and basal area from the yield table (
Table 1).
Table 1.
Stand data for initialization in 1951 and 1994, respectively (dg = mean tree diameter at breast height, hg = mean tree height).
Table 1.
Stand data for initialization in 1951 and 1994, respectively (dg = mean tree diameter at breast height, hg = mean tree height).
| | 1951 | 1994 |
---|
Age (years) | dg (cm) | hg (m) | Stock (m3) | Age (years) | dg (cm) | hg (m) | Stock (m3) |
---|
P1 | pine | 18 | 9.0 | 13.7 | 209 | 62 | 29.8 | 25.9 | 458 |
P2 | pine | 29 | 10.5 | 14.0 | 205 | 73 | 28.3 | 24.3 | 347 |
M1 | pine | 85 | 31.8 | 25.0 | 258 | | | | |
oak | 85 | 20.9 | 20.0 | 136 |
M2 | pine | 65 | 24.8 | 23.1 | 159 | | | | |
oak | 70 | 21.2 | 22.4 | 239 |
Soil data were available from soil inventories. The soil characteristics differ between the pure and the mixed stands. In particular, the sum of plant available water is much greater for the mixed stands (
Table 2).
Table 2.
Soil characteristics from soil inventory (WsFC = Sum field capacity, WsAW = Plant available water at maximum rooting depth, C/N = Carbon Nitrogen ratio total soil, ps = range sand fraction, pc = range clay fraction).
Table 2.
Soil characteristics from soil inventory (WsFC = Sum field capacity, WsAW = Plant available water at maximum rooting depth, C/N = Carbon Nitrogen ratio total soil, ps = range sand fraction, pc = range clay fraction).
Site | Soil Type | Soil Depth (cm) | WsFC (mm) | WsAW (mm) | C/N | ps (%) | pc (%) |
---|
P1 | brunic arenosol | 270 | 217 | 119 | 22 | 90–100 | 0–5 |
P2 | brunic arenosol | 255 | 237 | 93 | 21 | 50–100 | 0–17 |
M1 | brunic arenosol | 308 | 526 | 369 | 21 | 91–98 | 1–4 |
M2 | brunic arenosol | 305 | 607 | 395 | 15 | 75–97 | 2–9 |
2.4. Climate Data
The climate data are provided by climate stations, which are located nearby the forest stands. These climate data include daily measurements of weather parameters (temperature, precipitation, radiation, air humidity, vapor pressure, wind velocity) from 1951–2006 (Table S2). In the case of the pure pine stands (P1, P2), climate data of a mobile climate station next to the forest stand was also available from 1996–2004. For these stands, we merged the long time series of the permanent climate station with the short but more precise one of the mobile climate station wherever possible.
The climate conditions at the forest sites represent the subcontinental climate prevailing in Brandenburg with low precipitation in the growing season but slightly differ in precipitation and temperature (
Table 3).
Table 3.
Climatic characterization of forest sites on the base of permanent climate stations (Ty = yearly average temperature, Tg = growing season (April–September) average temperature, Py = average annual sum of precipitation, Pg = growing season (April–September) average sum of precipitation.
Table 3.
Climatic characterization of forest sites on the base of permanent climate stations (Ty = yearly average temperature, Tg = growing season (April–September) average temperature, Py = average annual sum of precipitation, Pg = growing season (April–September) average sum of precipitation.
Site | Ty (°C) | Tg (°C) | Py (mm) | Pg (mm) |
---|
P1 | 8.1 | 13.8 | 600 | 344 |
P2 | 8.5 | 14.7 | 550 | 309 |
M1 | 8.5 | 15.2 | 531 | 255 |
M2 | 8.9 | 16.2 | 521 | 267 |
2.5. Measured Data
2.5.1. Xylem Sap Flow Data
Tree canopy transpiration was estimated by sap flow measurements in 10 representative pine trees of stand P1 using the constant heating method based on Granier [
39]. The method is described in detail for this dataset by Lüttschwager
et al. [
40]. The stand sap flow was calculated as the product of average sap flow density and stand sapwood area that was estimated by stand inventory and sapwood area in the sample trees. The measurements were taken from June 1998 until October 1999. The aggregated stand transpiration estimated from the measured data was compared with simulated stand transpiration (
Table 4).
Table 4.
Summary of data and measurement periods available for each forest stand considered in this study.
Table 4.
Summary of data and measurement periods available for each forest stand considered in this study.
Site | Data | Period | Source |
---|
P1 | | 06/1998–10/1999 | [40] |
1997–2004 | [41,42,43] |
until 2004 | [44] |
P2 | | 1997–2004 | [41,42,43] |
until 2004 | [44] |
M1 | | until 2006 | [45] |
M2 | | until 2006 | [45] |
2.5.2. Soil Water Content
The volumetric soil water content was taken from [
41,
42,
43] and calculated using the time domain reflectometry (TDR) measurements at the P1 and P2 sites. The TDR tubes were installed in three different soil depths (P1: 20, 70, 250 cm and P2: 20, 50, 250 cm). The observation period lasted from 1997 until 2004 (
Table 4).
2.5.3. Tree Ring Data
The growth dynamics of the trees was measured on the base of annual ring widths taken from increment cores. In each case, two increment cores were extracted from 20 dominant pines and 20 dominant/subdominant oaks at the M1 and M2 sites in 2006 [
45]. At P1 and P2, only average annual ring width time series until 2004 for dominant pine trees were available [
44]. For the comparison between simulated and measured ring widths, we also calculated the average annual ring width time series at the mixed forest stands M1 and M2 (
Table 4).
2.6. Validation Procedure
The validation of the different model approaches to simulate water uptake were executed with different variables available from the different measurement periods. The processes that are important for tree growth are simulated at different time scales. The soil water balance is simulated with a daily time step and, consequently, the validation is also done with daily data. The short-term validation with the transpiration measurements on P1 encompasses one and a half years and the soil water measurements on P1 and P2 eight years. The annual diameter increment is simulated at the end of the year when the carbon is allocated to the stem. Therefore, the long-term validation is executed with annual data with the tree ring chronologies from 1951–2004 (P1, P2) and 1951–2006 (M1, M2), respectively.
The average annual ring increment for each species and site were used to derive index series through normalization. The index values were calculated in two steps. Firstly, an autoregressive model converts increment values into index values:
where
rwt is the ring width and
rwt−1 is the ring width of previous year,
a and
b are empirical parameters.
Secondly, a linear regression minimizes the tree age impacts:
where
tage is the tree age,
c and
d are empirical parameters.
We compared the index time series of simulated and observed diameter growth by considering four criteria. The first criterion is the
Pearson correlation coefficient
r (Equation (3)):
where
Xi is the
ith simulated value (
i = 1, ..,
n),
X̄ is the arithmetic mean of simulated values,
Yi is the
i-th measured value (
i = 1, ..,
n),
Ȳ is the arithmetic mean of measured values.
For the soil water content we used the coefficient of determination
R2 between simulated and measured values:
where
Ŷi is the
ith estimated value of the regression between simulated and measured values,
Ȳ the arithmetic mean of simulated values,
Yi the
ith simulated value and
n the number of pairs of simulated and measured values.
The sensitivity index (SI) indicates the year-to-year fluctuations of ring widths [
44]. Sensitivity index is calculated using the following equation:
The third criterion is the “Gleichläufigkeit” score
Gxy of two tree ring index series
xi and
yi (
i = 1, …,
n) and is defined by:
with:
This index counts how well the two series have followed each other over the years. The last criterion is a drought index (
DI) to detect marker years with small diameter growth due to water shortage. Following Schröder
et al. [
45], we defined a marker year as a year where the diameter increment of a year is at minimum 10% above or below the increment of the previous year. Thereafter, only the lower increments were used to filter out the drought years. Because complex interactions between climate, management and diameter growth can confound the effect of drought, we filtered out years where remarkably low diameter increment should mainly be caused by water shortage. For this filtering, a cluster analysis was performed on the base of the variables minimum water content in the soil, climatic water balance of the year and number of ice days (
Tmax < 0 °C). At each site, the classification of the marker years to three clusters was based on the Euclidean distance. As a result, marker years with dry conditions could be separated from marker years with a strong winter and marker years with wet conditions. The resulting drought years (Table S3) fit well to reported drought years in the northeastern lowlands of Germany. The cluster analysis was conducted with the software package “
cluster” available for
R [
46,
47]. The drought index,
DI, is then calculated as the percentage concurrence of observed and simulated drought years.
2.7. Statistical Analysis
To find an over all result of the performance of the water uptake approaches related to the annual diameter increment we aggregated the four criteria described above into one metric. Since all four criteria can be assessed with a scale from 0% to 100%, they were standardized to allow calculating a mean value for the water uptake approaches. Hereby, the standardization of the four criteria is as follows:
- criteria I
r* = r × 100 (if r < 0; r* = 0)
- criteria II
Gxy* = Gxy
- criteria III
SI* = 100 − (|SIsim − SImeas|)/SImeas × 100)
- criteria IV
DI* = DI
The asterisk indicates the standardized criterion and a value of 100% means perfect compliance with the criterion of the measured index time series.
The Tukey test (tukeyHSD, package
stats) [
47] was used to test for significant differences between the four average compliance values of the four water uptake approaches WU1, WU2-low, WU2-medium, and WU2-high. It is a nonparametric test for multiple comparisons. The performance of the water uptake approaches simulating stand tree transpiration and soil water content at a specific soil depth was evaluated using the coefficient of determination
R2 between simulated and measured values (Equation (11)). The coefficient of variation (
CV) as the ratio between the standard deviation and the mean value was calculated to compare variation in simulated and observed soil water content. The statistical analyses were conducted with equations implemented in 4C and with the package
“base” statistical software
R version 2.10.0 [
47].
5. Conclusions
This paper integrated two water uptake approaches—an empirical one and a process-based one—in a process-based forest growth model and evaluated their performance in Scots pine and Scots pine-Sessile oak stands in Germany. We firstly assessed whether there are main differences between the two water uptake approaches regarding the water supply from the roots. We show that the main difference between them is the nonlinear relationship between soil water content and soil water potential, which determines the soil water resistance. This induces different root water uptake in the process-based approach in comparison to the empirical one, which is linearly dependent on a soil water content reduction factor. The study also showed the high influence of the assumed total root resistance on water uptake. In the empirical water uptake approach, the modeled root distribution and, consequently, root length densities in different soil layer have negligible effects on water uptake. In contrast, the modeled root length densities lead to high water uptake limitations in the process-based approach with high total root resistance.
Secondly, we analyzed how well drought years could be reproduced by the different water uptake approaches. We found the highest compliance between observed and simulated drought years in the process-based water uptake approach with high total root resistance.
Finally, we assessed whether there was one approach that performed best regarding the different validation data sets. This study shows that there is not one approach that performs best regarding all three validation data sets. For a better validation of both approaches, it would be necessary to include measurements of root length densities and water uptake rates of fine roots on the same site. However, despite these further data requirements for more advanced model evaluation, this study clearly indicates that forest process-based models aiming at simulating drought conditions require further improvement in terms of model formulations and validation of different processes at various time scales. Further, we highlight the importance of a realistic modeling of understorey vegetation transpiration, especially in mature pine stands with high understorey vegetation cover.