Abstract
The within-species diversity in response to weather and the gaps in the response diversity in the modern set of forage crop cultivars were determined using an approach that assessed the adaptive capacity under global climate change. The annual dry matter (DM) yields were recorded in multi-location MTT (Maa- ja elintarviketalouden tutkimuskeskus) Agrifood Research Official Variety Trials in Finland for modern forage crop cultivars from 2000 to 2012, as a response to agroclimatic variables critical to yield based on the year-round weather data. The effect and interaction of cultivars and agroclimatic variables were analysed using mixed model. The relatively low adaptive capacity of timothy (Phleum pratense L.) and meadow fescue (Festuca pratensis Huds.) indicates that diversification of the breeding material is warranted, particularly for resistance to high temperatures during primary growth and to high temperature sum 7 days after the first cut. All red clover cultivars (Trifolium pratense L.) suffered from both low and high accumulation of warm winter temperatures. Except for the red clover cultivars, cold stress during winter and lack of warm winter temperatures consistently reduced the yields of all species and cultivars. All tall fescue (Festuca arundinacea Schreb.) cultivars suffered from low precipitation during the fall hardening period. Although the set of festulolium (Festulolium pabulare) cultivars was also sensitive to low precipitation during the fall, festulolium was a good example of enhanced capacity to adapt to climate change with high response diversity because the cultivar germplasm base was diversified. Foreign origin in a cultivar pool was apparently not sufficient or necessary to ensure added value for a diversity of responses to climate change. Similar analyses to those used in this study, applied as practical tools for breeders, farmers and public actors, are important to secure the adaptive capacity of crops worldwide under global climate change.
Similar content being viewed by others
Avoid common mistakes on your manuscript.
1 Introduction
Adaptation of crops to global climate change represents an urgent challenge for humanity. Although crops can be relatively well adapted to slow changes in the climate, the capacity to adapt to increased seasonal and inter-annual variation and to increased frequency of extremes may not be as high (Porter and Semenov 2005). Such potential vulnerability of agricultural production has created interest in building adaptive capacity and resilience (Smit and Wandel 2006; Lin 2011).
Dairy farming is the agricultural production line of highest economic significance in northern Europe, which is experiencing the most rapid climate change. Forage crops are the cornerstones in such production systems. In the future, forage production could benefit from several effects of climate change (Bindi and Olesen 2011; Graux et al. 2013; Rapacz et al. 2014), which include a longer physiologically effective growing season, a higher temperature sum and a higher concentration of CO2 (Porter et al. 2014) and better overwintering conditions that are predicted for some winter-sensitive species (Thorsen and Höglind 2010). As threats to the maintenance of sustainable forage production, increased frequency (Christidis et al. 2014) and duration of heat stress, droughts and heavy rain events are likely (Christensen and Christensen 2007; Ylhäisi et al. 2010; Ruosteenoja et al. 2011). Furthermore, climate change is expected to increase the risks of winter injury for most overwintering forages (Bélanger et al. 2002; Bertrand et al. 2003; Thorsen and Höglind 2010). A lack of full hardening during the fall and reduced snow cover may also expose crops to killing frosts, ice encasement and soil heaving (Bélanger et al. 2002; Rapacz et al. 2014).
In the context of climate change, ‘adaptive capacity is the ability of a system to adjust to climate change, including climate variability and extremes, to moderate potential damages, to take advantage of opportunities, or to cope with the consequences’ (IPCC 2001). Therefore, high adaptation is different from high adaptive capacity (Gallopín 2006; Smit and Wandel 2006), and a species or cultivar may be well adapted to the current environment but have little capacity to respond to variability and changes. We are unable to determine which cultivars will perform well in a future climate, but we may assume the requirements for capacity to adapt are likely to broaden (Coumou and Rahmstorf 2012; Chen and Tian 2016). Consequently, sufficient plasticity can hardly be bred within a single cultivar but rather within the entire cultivar pool. A key complement of the ability to respond is response diversity, meaning the diversity of responses within a functional group (Chapin et al. 1997; Elmqvist et al. 2003; Kahiluoto et al. 2014). Such diversity of responses to critical environmental change is likely to have a value that is proportional to its amount (Kahiluoto et al. 2014). Diversity of within-species responses in the available cultivar pool could increase yield stability under the current level of weather variability and ensure the potential for adaptation in the long-term as changes in the global climate proceed.
The aim of the study was to illustrate that assessments of response diversity can be used to identify gaps in the adaptive capacity to climate change. Specifically, the aim was to investigate whether within-species diversity in responses was apparent to changing and varying weather for the available set of modern forage crop cultivars as the most important crops grown in northern Europe: timothy (Phleum pratense L.), meadow fescue (Festuca pratensis Huds.), tall fescue (Festuca arundinacea Schreb.; syn. Lolium arundinaceum Schreb), tall fescue-type festulolium (F. arundinacea Schreb. × Lolium multiflorum Lam, i.e., Festulolium pabulare) that were backcrossed with F. arundinacea (DLF 2016) and red clover (Trifolium pratense L.). The annual forage crop dry matter (DM) yield (kg ha−1 year−1) was used throughout the study as the response variable to indicate adaptive capacity to agroclimatic variables critical to yield. The specific research questions were as follows:
-
1.
Is there within-species diversity in responses to critical agroclimatic variables in the available set of modern forage crop cultivars?
-
2.
To enhance the adaptive capacity, to which agroclimatic variables are the diversity of responses low?
2 Materials and methods
2.1 Variety trials
The Official Variety Trial results from MTT Agrifood Research Finland were used (Kangas et al. 2009). The Official Variety Trial data consisted of 8361 yield records for 126 cultivars from 16 different trial sites from 1980 to 2012. In this study, modern cultivars from 2000 to 2012 were selected from the following species: timothy, meadow fescue, tall fescue, festulolium and red clover. There were 1156 records of annual DM yields (kg DM ha−1 year−1) for 39 modern cultivars (Table 1). However, the entire data set from 1980 to 2012 was used in the analyses because every year the set of the tested cultivars changed slightly and thus long-term control cultivars improve the accuracy of DM yield estimates of modern cultivars. Annual DM yields (kg DM ha−1 year−1) were used as a response variable because DM yield can be used as a proxy for variation in food supply and farm income, and yield is also an indicator of ecosystem health (Rapport et al. 1998). Cultivars were of Finnish and foreign origin (Online resource 1), and each cultivar was associated with 20 to 70 observations from 11 different trial sites. All experiments at all sites were arranged as randomized complete block designs or incomplete block designs with three or four replicates. Plot size was 7–10 m × 1·25 m, depending on location and year. The experimental design and the management were similar to that on-farm, as described in more detail by Kangas et al. (2009) and Hakala et al. (2012).
The forage stands were productive 3 (to four) years after the establishment year and were harvested two to three times per year. The harvest dates varied among years and trials because of weather conditions and trial locations (e.g., latitude and soil type). A few harvest dates were missing (14 of 1409), which were estimated using a linear mixed model, relying on data for harvest dates of other cultivars at the same site and year. The effects of the site, year and species were considered in the model. The trials with missing data for the growing season were not included in the estimation of the variables. A flow diagram (Fig. 1) describes the steps of the following analysis.
2.2 Selecting the critical factors of change and variation
The year-round daily weather data (1979–2012) from the Finnish Meteorological Institute for weather stations closest to the Official Variety Trials were used to assess the yield response to agroclimatic variables for each species and their cultivars separately. We identified agroclimatic variables critical for the forage yield performance in northern Europe using linear mixed models (Mäkinen et al. 2015); the yield responses were calculated for 35 pre-selected agroclimatic variables, and the agroclimatic variables were removed that, among other reasons, had a nonsignificant effect, minor effect or were missing too much information. In this study, these previously suggested critical agroclimatic variables (for details, see Mäkinen et al. 2015) were tested regarding the within-species diversity of a modern set of forage crop cultivars (Table 2).
The agroclimatic variables were preliminarily selected based on published literature (Bélanger et al. 2002; Volenec and Nelson 2007; Thorsen and Höglind 2010). The fall hardening period (FH), winter period (WP) and growth period (GP) were distinguished as suggested by Thorsen and Höglind (2010) and Bélanger et al. (2002). Regarding the FH and WP, the agroclimatic variables suggested by Thorsen and Höglind (2010) and Bélanger et al. (2002) were used. Different temperatures and precipitation rates were tested during the GP, based on the optimum temperatures for the cool season grasses suggested by Volenec and Nelson (2007). Because the development of forage crops is faster and characterized by different tiller traits during the primary growth stage, in comparison with the regrowth stage (Bélanger and McQueen 1998; Virkajärvi et al. 2012), some of the temperature and precipitation variables were tested separately for the first and the second harvest.
2.3 Categorizing agroclimatic variables
Agroclimatic variables were analysed separately to investigate the gaps in the adaptive capacity of cultivars. Some agroclimatic variables were also strongly correlated leading to multicollinearity in regression analysis. The assumptions of linearity were also violated because the relations between the yield and the agroclimatic variables were nonlinear in most cases. Most importantly, we had to consider that the random effects of year, site and experiment contained most of the total variation; therefore, linear mixed models were used in data analyses.
Each agroclimatic variable was divided into three categories (low, moderate and high; Table 2) (Hakala et al. 2012). Trial observations were divided equally into the three categories to ensure the reliability of yield estimates. For example, the number of cold stress days during the winter period (W-STRESS) was in the low category for 0–10 days, in the moderate category for 11–21 days and in the high category for more than 21 days. We used a minimum requirement of 20 observations per cultivar together with possible missing values in a random agro-climatic variable. More uneven division of observations would have led to categories with few observations only for some agroclimatic variables.
2.4 Estimating responses to the factors
The main effects and interaction of cultivars (G = genotype) and categorical agroclimatic variable (E = environment) were analysed for each species separately using the following mixed model:
where y ijklm is the observed yield (annual DM yield), μ is the intercept, cultivar i is the average yield level of the ith cultivar, category j is the average yield level at the jth level of categorized environment (j = 1, 2, 3) and cultivar × category ij is the cultivar-by-environment interaction. All of the above effects are fixed in the model. Experimental site × year × trial(category) jklm is the random effect of the kth experimental site, lth year and mth trial within the jth category and ε ijklm is a normally distributed residual error.
2.5 Looking for gaps in the yield responses of species
P values of statistical significance for G, E and G × E were calculated. Although a clear response was indicated for cases in which E was statistically significant but G × E was not, the response diversity within the cultivars of a species was low in these cases, and therefore, a gap in adaptive capacity was identified. The results are illustrated in figures regarding those agroclimatic variables with the most diversity in responses and those with the least (Figs. 2, 3, 4, 5 and 6 and Table 4).
Statistical analyses were performed using the MIXED procedure in the SAS statistical software package (version 9.3, SAS Institute Inc., Cary, NC, USA).
3 Results
3.1 Timothy
Modern timothy cultivars showed no differences in yield responses to the agroclimatic variables (Table 3). The cultivars suffered from high precipitation during FH (Fig. 2a), and a long FH decreased the yields by more than 5 % in all cultivars (Table 4).
In the growth period, all cultivars were sensitive to the high accumulation of days with temperature of 25 °C during the primary growth (Fig. 2b) (see Online resource 2).
3.2 Meadow fescue
Meadow fescue cultivars responded differently to one of the 14 agroclimatic variables tested (Table 3). All cultivars showed sensitivity to low precipitation during the fall hardening period (Fig. 3a). Additionally, the occurrence of a high number of cold winter temperatures (>−15 °C) consistently reduced the yields by 10–25 %. A high accumulation of warm winter temperatures (above 0 °C) increased the yields nearly 20 % compared with those of the lowest temperature accumulation category (Table 4).
Regarding the growth period, diversity occurred in the yield responses to a high number of days with the maximum temperature of 25 °C during the regrowth stage (Fig. 3b); in most cases, the temperature stress led to marked yield reduction, but SW Minto and Valtteri cultivars yielded well under the moderate and high rates of temperature accumulation. A high temperature sum 7 days after the first cut reduced the yields of all cultivars (Fig. 3c) (see Online resource 3).
3.3 Tall fescue
Tall fescue cultivars showed diversity in yield responses to two of the 14 agroclimatic variables (Table 3). During fall hardening, an increase in accumulation of cold temperatures <5 °C consistently reduced the yields, and in the lowest precipitation category, yield penalties occurred in all tall fescue cultivars (Fig. 4a). All tall fescue cultivars were apparently sensitive also to harsh winter conditions: yield reductions of approximately 15 % were found during winters characterized with a high number of days causing cold stress (−15 °C; Table 4). A high accumulation of warm winter temperatures (above 0 °C) increased the yields by approximately 20 % compared with those of the lowest accumulation category.
In the growth period, tall fescue cultivars differed in response to the temperature sum accumulation and to the number of days with the maximum temperature of 28 °C. However, a high temperature sum 7 days after the first cut was not tolerated well by any of the cultivars (Fig. 4b) (see Online resource 4).
3.4 Festulolium
Festulolium cultivars showed differences in yield responses to eight of the 14 agroclimatic variables (Table 3). All cultivars were apparently sensitive to a low or moderate rate of precipitation during the fall hardening period (Fig. 5a). All festulolium cultivars were sensitive to the high accumulation of cold temperatures (−15 °C) during the winter period (10–21 % yield reduction). By contrast, a high accumulation of temperatures above 0 °C during the winter period increased yields markedly (Table 4).
During the growth period, the cultivars differed in their yield responses to the temperature sum. Cultivars differed in their responses to the number of days with the maximum temperature of 25 °C in the primary and regrowth stages (Fig. 5b). Cultivars also differed in their responses to very high temperature stress (28 °C) during the growth period. In most cases, the accumulation of a high temperature sum 7 days after the first cut led to a yield reduction. Cultivars responded differently to the accumulation of precipitation before the first cut (see Online resource 5).
3.5 Red clover
The modern red clover cultivars showed differences in yield responses to one of the 14 agroclimatic variables and a tendency in responses to three of the 14 (Table 3). Although red clover cultivars were sensitive to winter temperatures that differed from the moderate accumulation of temperatures above 0 °C, there was a tendency towards differences in yield responses (Fig. 6a).
The highest (and the lowest) number of days of the very high temperature stress (+28 °C) led consistently to a yield reduction (Fig. 6b). The low temperature accumulation rate or the temperature sum led to marked yield reductions (Table 4) (see Online resource 6).
4 Discussion
4.1 Sensitivity of forage crop cultivars to climate change
We used the annual forage crop DM yield (kg ha−1 year−1) to indicate the response diversity as a proxy to adaptive capacity to weather variability throughout the study. Although the DM yield of the first cut is typically more sensitive than the annual harvest to changes in the fall hardening and winter related variables (Kangas et al. 2009), the annual DM yield is by far the most important variable for forage production because it is linked to farm fodder and food supply and farm income.
The length of the hardening period is associated with better winter survival of grasses (Bélanger et al. 2002). According to Bélanger et al. (2002), the mean length of fall hardening in eastern Canada is approximately 24 days, whereas in this study, the mean length of lowest category of the fall hardening period was 39 days (Table 2). This seems sufficiently long for the tested forage crop cultivars, since it did not result in marked damage. If precipitation during the fall increases as projected (Ruosteenoja et al. 2011), excessive soil moisture is likely to hamper the hardening (Paquin and Mehuys 1980), as indicated by our findings regarding yield penalties in the timothy cultivars, whereas other forage crop cultivars succeeded well under such conditions. In this regard, diversity in the breeding material of timothy is required to complement the new drainage measures.
According to our results, warm winter temperatures increased the yields of timothy, meadow fescue, festulolium and tall fescue cultivars, as also reported by Thorsen and Höglind (2010), wherein the occurrence of ice cover is rarely reported. Höglind et al. (2010) found that timothy cv. Grindstad was more sensitive to frost than cv. Engmo, but we did not establish differences among the timothy cultivars in responses to those tested winter weather variables. All modern red clover cultivars succeeded well under the moderate accumulation of warm winter temperatures. The sensitivity of red clover cultivars to the high accumulation of warm winter temperatures might be explained by dehardening and consequent winter damage (Bélanger et al. 2002). On the other hand, the sensitivity of the cultivars to the low accumulation of such warm winter temperatures and associated long-standing snow cover might be explained by the promotion of diseases such as clover rot (Sclerotinia trifoliorum) (Ylimäki 1967).
The expected elevated temperatures during the growing season combined with long days may disturb growth and result in overly rapid crop development in the future. Temperature stress reduces the growth and chlorophyll content and causes injuries to cell membranes (Heide 1982; Wang et al. 2009). Temperatures below 21 °C, particularly at approximately 17 °C, are suggested for the optimal growth of timothy (Smith 1972; Bertrand et al. 2008), which explained the sensitivity of timothy cultivars to temperature stress observed in our study. Unlike our findings for DM yields of timothy cultivars under field conditions, significant differences are reported for temperature tolerance in timothy cultivars in growth chamber assays (Heide 1982). For example, high temperatures apparently more easily inhibit the flowering of cultivars of Scandinavian origin than that of British or American cultivars (Cooper 1958). Whereas temperature stress was harmful for most of the tested meadow fescue cultivars, the cultivar Valtteri, which is of Finnish origin, performed better under hot conditions. Regarding tall fescue, studies show great variability among the cultivars in responses to temperature stress (Wang et al. 2009; Sun et al. 2015), such as we found for both tall fescue and festulolium. Red clover cultivars, as reported earlier (Fagerberg 1988), were not sensitive to the occurrence of days with the maximum temperature of 25 °C. However, we did show a marked yield reduction in the red clover cultivars when exposed to a high number of days with the maximum temperature of 28 °C. A high temperature sum 7 days after the first cut consistently reduced yields in all of the meadow fescue and tall fescue cultivars tested.
Climate change is expected to change patterns of precipitation, and in particular, early season droughts may limit the yield potential in Finland (Trnka et al. 2011). Although timothy has high root biomass (Bolinder et al. 2002), it has low tolerance for drought (Molyneux and Davies 1983; Wilman et al. 1998). However, no significant yield penalties for the timothy cultivars were shown for conditions of low precipitation in the primary growth and regrowth stages. The high soil water reserves in spring due to melting snow might have contributed to these results. The persistence of most of the tested fescue species to summer droughts was previously reported (Humphreys et al. 1997). A variation in root systems among genotypes, which has been reported, e.g. in a perennial ryegrass (Crush et al. 2007), might have affected our findings for festulolium, because yield responses of festulolium cultivars differed greatly according to different precipitation rates during the primary growth stage. We did not find any marked effect of precipitation-related weather variables on tall fescue and meadow fescue yields or on differences among the responses of cultivars, although Huang and Gao (1999) and Assuero et al. (2002) previously reported high levels of variation among tall fescue cultivars. All red clover cultivars suffered from dry conditions during regrowth, as also reported by Mela (2008) and Fagerberg (1988).
4.2 Diversity of responses in the modern set of forage crop cultivars
Overall, when compared with barley cultivars (Hakala et al. 2012), the modern forage crop cultivars responded more similarly to the tested agroclimatic variables. The high diversity observed in the responses of festulolium cultivars was likely due to the diverse genetic base, because the festulolium cultivars were synthetic F. arundinacea × L. multiflorum hybrids (DLF 2016). By contrast, the similarity of responses observed for the timothy cultivars was likely due to the cross-pollinating that occurs with timothy. According to Tanhuanpää and Manninen (2012), a lack of differentiation in Nordic timothy is likely due to the substantial gene flow between cultivars and natural populations, which prevents geographical differentiation. However, they concluded that the timothy material apparently has sufficient diversity for breeding purposes (Tanhuanpää and Manninen 2012), although genetic variation for stress adaptation was not considered. In the set of meadow fescue cultivars from Nordic countries, most of the genetic variation occurs within populations rather than among different cultivars (Fjellheim and Rognli 2005), which explained our findings for meadow fescue. Considerably low levels of genetic variation are found for meadow fescue cultivars compared with perennial ryegrass or cocksfoot cultivars (Kölliker et al. 1999), and such a limited genetic variability likely contributed to the decline in the use of meadow fescue for grass production (Kölliker et al. 1999). Although we found significant differences in yield responses of the modern cultivars to some weather variables, we assume that variation will likely increase when the breeding includes a wider range of red clover (e.g. Helgadóttir 1996; Halling et al. 2004) and tall fescue (e.g. Mian et al. 2002; Majidi et al. 2009) cultivars.
4.3 Towards enhanced adaptive capacity of forage crops
Effective adaptation is likely not possible without breeding programs (Rosenzweig and Tubiello 2007). Investigations of the response diversity by cultivars of a broader geographical origin and natural populations, in addition to locally adapted populations, such as those reported for meadow fescue (Kanapeckas et al. 2005), would be very valuable in breeding to build the adaptive capacity of forage crops. Instead of increasing only the genetic diversity of a cultivar pool, the identification of sources of response diversity to climate change ensures diversity that is more effective for adaptation of forage crops to climate change (Chapin et al. 1997; Elmqvist et al. 2003; Kahiluoto et al. 2014). Even if genetic diversity may not enhance response diversity per se (Mäkinen et al. 2015), Festulolium cultivars (synthetic F. arundinacea × L. multiflorum hybrids) exemplify the significance of a broad genetic basis for building adaptive capacity within crops (Reed and Frankham 2003; Ehlers et al. 2008). However, this type of synthetic species hybridization is most likely impractical for many other grass species. It was observed that cultivars of foreign origin (such as the timothy cultivars Lischa and Moverdi and Suez red clover) tended to respond differently when compared with the cultivars of Finnish origin. However, foreign origin inclusion was not sufficient to achieve diversity in response, as indicated by the example of the heat-resistant meadow Finnish fescue cultivar Valtteri. Our forage crop-specific results are applicable to northern European conditions, for example, by cultivating cultivars that respond differently in different fields of one farm or of one region (Himanen et al. 2013; Mäkinen et al. 2015) or by using mixtures of cultivars within the same field (Kiær et al. 2009; Tooker and Frank 2012). Further investigations that include the sensitivity of the digestibility of the forage yield to variability in critical weather components are also essential (Thorvaldsson et al. 2007; Bertrand et al. 2008).
4.4 Global relevance
Under the intensified variability, extremes (Coumou and Rahmstorf 2012) and uncertainty (Rötter et al. 2013) of global climate change, strategies for building capacity to adapt to various plausible changes are required. Such capacity-oriented strategies complement the globally prevalent strategies to ‘predict-and-adapt’ to long-term average changes in climate that are difficult to predict (Dessai et al. 2007) as highlighted by our results. The proposed approach to assess and increase the response diversity represents a strategy to safeguard adaptation of crops without precise climate predictions, acknowledging uncertainty. Such adaptation strategies are important to apply more generally to food and feed crops that are cornerstones of global and national food security (Trnka et al. 2014). Through similar assessments as exemplified here, potential gaps in the adaptive capacity can be recognized and bridged. This is particularly important given the inherent vulnerability of the global food supply that has become relatively species-poor during the past 50 years (Khourya et al. 2014) and also nationally homogeneous worldwide.
Currently, the adaptive capacity of commercial crops is largely determined by private plant breeding companies in most regions around the world. Thus, because of the highly competitive market with a high cost pressure, the size of the cultivar market may determine the breeding goals. Consequently, some traits such as sensitivity to a particularly important weather variable might go unnoticed, which could contribute to a gap in the adaptive capacity of the cultivar pool. Specific tools for increasing the response diversity within species through breeding and diversification of cultivated crops will require further studies and the involvement of the relevant stakeholders, including both breeders and farmers (Paavola et al. 2016). Public actors, such as national emergency supply agencies or relevant ministries, should be included in setting the cultivation and breeding goals and financing breeding for the adaptive capacity in crops to provide stable food and fodder, because longer term goals may not be privately financed in a competitive market.
Regarding the presumably more frequent and intensive extreme conditions in the future (Christidis et al. 2014) anywhere around the world, an option to apply the assessment illustrated here would be to use within each weather variable a wider moderate category (such as 50 %) in the analysis, while here, each weather category (low, moderate and high) included a third of the observations. Such a modified assessment would give more weight to the capacity to adapt to extreme conditions in the response diversity assessment.
5 Conclusions
Cultivars should be developed for adaptation to the variability of and extremes in weather and for multiple scenarios for mid- and long-term perspectives for global climate change. In the present study, empirical assessments of the sensitivity of yield of a modern set of forage cultivars to a broad range of critical weather conditions revealed the adaptive capacity of the cultivar pool and also its weak points; thus, such empirical assessments can be a practical tool for breeding for an uncertain future. Whereas the low adaptive capacity of timothy and meadow fescue indicate the development of more diverse breeding material is warranted, the set of festulolium cultivars was a good example of enhanced adaptive capacity to climate change through diversification of the cultivar germplasm base, even if synthetic species hybridization such as taken place for festulolium is most likely impractical for many other grass species. Furthermore, the inclusion of foreign origin in a cultivar pool was not necessary to ensure added value for response diversity. Similar analyses as exemplified here are useful regarding the entire set of food and feed crops, to secure the capacity to adapt to the variability and changes driven by global climate change. Diversity in responses to weather as one example of using within-species variation to increase the capacity to adapt to global climate change (Carpenter et al. 2015) can contribute to an increase in food security. Therefore, practical tools based on similar empirical assessments are required for breeders, farmers and public actors, as well as for practitioners on many other fields around the globe.
References
Assuero SG, Matthew C, Kemp P et al (2002) Effects of water deficit on Mediterranean and temperate cultivars of tall fescue. Crop and Pasture Sci 53:29–40
Bélanger G, McQueen R (1998) Analysis of the nutritive value of timothy grown with varying N nutrition. Grass Forage Sci 53:109–119
Bélanger G, Rochette P, Castonguay Y et al (2002) Climate change and winter survival of perennial forage crops in eastern Canada. Agron J 95:1120–1130
Bertrand A, Tremblay G, Pelletier S et al (2008) Yield and nutritive value of timothy as affected by temperature, photoperiod and time of harvest. Grass Forage Sci 63:421–432
Bertrand A, Castonguay Y, Nadeau P et al (2003) Oxygen deficiency affects carbohydrate reserves in overwintering forage crops. J Exp Bot 54:1721–1730
Bindi M, Olesen J (2011) The responses of agriculture in Europe to climate change. Reg Env Change 11:151–158
Bolinder M, Angers D, Bélanger G et al (2002) Root biomass and shoot to root ratios of perennial forage crops in eastern Canada. Can J Plant Sci 82:731–737
Carpenter SR, Brock WA, Folke C et al (2015) Allowing variance may enlarge the safe operating space for exploited ecosystems. PNAS 112:14384–14389
Chapin FS, Walker BH, Hobbs RJ et al (1997) Biotic control over the functioning of ecosystems. Science 277:500–504
Chen X, Tian G (2016) Impacts of weather variations on rice yields in China based on province-level data. Reg Env Change 16:2155–2162
Christensen JH, Christensen OB (2007) A summary of the PRUDENCE model projections of changes in European climate by the end of this century. Clim Chang 81:7–30
Christidis N, Jones GS, Stott PA (2014) Dramatically increasing chance of extremely hot summers since the 2003 European heatwave. Nat Clim Chang 5:46–50
Cooper JP (1958). The effect of temperature and photoperiod on Flowering and reproduction in temperate grasses inflorescencedevelopment in strains of timothy (Phleum spp.). J Br Grassl Soc 13: 8l–91
Coumou D, Rahmstorf S (2012) A decade of weather extremes. Nat Clim Chang 2:491–496
Crush J, Easton H, Waller J et al (2007) Genotypic variation in patterns of root distribution, nitrate interception and response to moisture stress of a perennial ryegrass (Lolium perenne L.) mapping population. Grass Forage Sci 62:265–273
Dessai S, O’Brien K, Hulme M (2007) Editorial: on uncertainty and climate change. Glob Environ Chang 17:1–3
Dlf (2016) Forage seeds white paper. Festulolium hybrid grass, 4 p. Available at http://www.dlfis.com/upload/fesulolium_white_paper.pdf. Accessed 5 February 2016
Ehlers A, Worm B, Reusch TB (2008) Importance of genetic diversity in eelgrass Zostera marina for its resilience to global warming. Mar Ecol Prog Ser 355:1–7
Elmqvist T, Folke C, Nyström M et al (2003) Response diversity, ecosystem change, and resilience. Front Ecol Environ 1:488–494
Fagerberg B (1988) The change in nutritive value in timothy, red clover and lucerne in relation to phenological stage, cutting time and weather conditions. Acta Agric Scand 38:347–362
Fjellheim S, Rognli O (2005) Genetic diversity within and among Nordic meadow fescue (Huds.) cultivars determined on the basis of AFLP markers. Crop Sci 38:347–362
Fowler C, Hodgkin T (2004) Plant genetic resources for food and agriculture: assessing global availability. Annu Rev Environ Resour 29(1):143–179
Gallopín GC (2006) Linkages between vulnerability, resilience, and adaptive capacity. Glob Environ Chang 16:293–303
Graux A, Bellocchi G, Lardy R et al (2013) Ensemble modelling of climate change risks and opportunities for managed grasslands in France. Agric For Meteorol 170:114–131
Hakala K, Jauhiainen L, Himanen S et al (2012) Sensitivity of barley varieties to weather in Finland. J Agric Sci 150:145–160
Halling M, Topp C, Doyle C (2004) Aspects of the productivity of forage legumes in Northern Europe. Grass Forage Sci 59:331–344
Heide OM (1982) Effects of photoperiod and temperature on growth and flowering in Norwegian and British timothy cultivars (Phleum pratense L.). Acta Agric Scand 32:241–252
Helgadóttir Á (1996) Red clover (Trifolium pratense L.) varieties for northern regions. Acta Agric Scand Sec B 46:218–223
Himanen SJ, Ketoja E, Hakala K et al (2013) Cultivar diversity has great potential to increase yield of feed barley. Agron Sustain Dev 33:519–530
Höglind M, Bakken A, Jørgensen M et al (2010) Tolerance to frost and ice encasement in cultivars of timothy and perennial ryegrass during winter. Grass Forage Sci 65:431–445
Huang B, Gao H (1999) Physiological responses of diverse tall fescue cultivars to drought stress. Hortscience 34:897–901
Humphreys M, Thomas H, Harper J et al (1997) Dissecting drought-and cold-tolerance traits in the Lolium–Festuca complex by introgression mapping. New Phytol 137:55–60
IPCC (Intergovernmental Panel on Climate Change) (2001) Technical summary: climate change 2001: impacts, adaptation, and vulnerability. A Report of Working Group II of the Intergovernmental Panel on Climate Change
Kahiluoto H, Kaseva J, Hakala K et al (2014) Cultivating resilience by empirically revealing response diversity. Glob Environ Chang 25:186–193
Kanapeckas J, Tarakanovas P, Lemeþiene A (2005) Variability, heritability and correlations of genetic resources in meadow fescue. Biologija 3:10–14
Kangas A, Laine A, Niskanen M et al. (2009) Virallisten Lajikekokeiden Tulokset (Results of Official Variety Trials) 2002–2009. http://www.mtt.fi/mttkasvu/pdf/mttkasvu6.pdf
Khourya CK, Bjorkmanc AD, Dempewolfd H et al (2014) Increasing homogeneity in global food supplies and the implications for food security. PNAS 11:4001–4006
Kiær LP, Skovgaard IM, Østergård H (2009) Grain yield increase in cereal variety mixtures: a meta-analysis of field trials. Field Crops Res 114:361–373
Kölliker R, Stadelmann F, Reidy B et al (1999) Genetic variability of forage grass cultivars: a comparison of Festuca pratensis Huds., Lolium perenne L., and Dactylis glomerata L. Euphytica 106:261–270
Lin BB (2011) Resilience in agriculture through crop diversification: adaptive management for environmental change. Bioscience 61:183–193
Majidi MM, Mirlohi A, Amini F (2009) Genetic variation, heritability and correlations of agro-morphological traits in tall fescue (Festuca arundinacea Schreb.). Euphytica 167:323–331
Mäkinen H, Kaseva J, Virkajärvi P et al (2015) Managing resilience of forage crops to climate change through response diversity. Field Crops Res 183:23–30
Mela T (2008) Red clover grown in a mixture with grasses: yield, persistence and dynamics of quality characteristics. Agric Food Sci 12:195–212
Mian M, Hopkins AA, Zwonitzer JC (2002) Determination of genetic diversity in tall fescue with AFLP markers. Crop Sci 42:944–950
Molyneux D, Davies W (1983) Rooting pattern and water relations of three pasture grasses growing in drying soil. Oecologia 58:220–224
Paavola S, Himanen SJ, Kahiluoto H et al. (2016) Making sense of resilience in barley breeding: converting the concept of response diversity into a tool of reflection and decision-making. In: Paloviita A, Järvelä M (eds) Climate change adaptation and food supply chain management, pp. 43–54
Paquin R, Mehuys GR (1980) Influence of soil moisture on cold tolerance of alfalfa. Can J Plant Sci 60:139–147
Porter JR, Semenov MA (2005) Crop responses to climatic variation. Philosophical Transactions of the Royal Society B: Biological Sciences 360:2021–2035
Porter JR, Xie L, Challinor AJ et al (2014) Food security and food production systems. In: Field CB, Barros VR, Dokken DJ, Mach KJ, Mastrandrea MD (eds) Climate change 2014: impacts, adaptation, and vulnerability. Part A: global and sectoral aspects. Contribution of Working Group II to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, pp. 485–533
Rapport D, Costanza R, McMichael A (1998) Assessing ecosystem health. Trends Ecol Evol 13:397–402
Rapacz M, Ergon Å, Höglind M et al (2014) Overwintering of herbaceous plants in a changing climate. Still more questions than answers. Plant Sci 225:34–44
Rosenzweig C, Tubiello FN (2007) Adaptation and mitigation strategies in agriculture: an analysis of potential synergies. Mitig adapt strategies glob chang 12:855–873
Reed DH, Frankham R (2003) Correlation between fitness and genetic diversity. Conserv Biol 17:230–237
Ruosteenoja K, Räisänen J, Pirinen P (2011) Projected changes in thermal seasons and the growing season in Finland. Int J Climatol 31:1473–1487
Rötter RP, Höhn J, Trnka M et al (2013) Modelling shifts in agroclimate and crop cultivar response under climate change. Ecol Evol 3:4197–4214
Smit B, Wandel J (2006) Adaptation, adaptive capacity and vulnerability. Glob Environ Chang 16:282–292
Smith D (1972) Effect of day-night temperature regimes on growth and morphological development of timothy plants derived from winter and summer tillers. Grass Forage Sci 27:107–110
Sun X, Du Z, Ren J et al (2015) Association of SSR markers with functional traits from heat stress in diverse tall fescue accessions. BMC Plant Biol 15:116
Tanhuanpää P, Manninen O (2012) High SSR diversity but little differentiation between accessions of Nordic timothy (Phleum pratense L.). Hereditas 149:114–127
Thorsen SM, Höglind M (2010) Assessing winter survival of forage grasses in Norway under future climate scenarios by simulating potential frost tolerance in combination with simple agroclimatic indices. Agric For Meteorol 150:1272–1282
Thorvaldsson G, Tremblay GF, Tapani Kunelius H (2007) The effects of growth temperature on digestibility and fibre concentration of seven temperate grass species. Acta Agric Scand 57:322–328
Tooker JF, Frank SD (2012) Genotypically diverse cultivar mixtures for insect pest management and increased crop yields. J Appl Ecol 49:974–985
Trnka M, Olesen JE, Kersebaum KC et al (2011) Agroclimatic conditions in Europe under climate change. Glob Chang Biol 17:2298–2318
Trnka M, Rötter RP, Ruiz-Ramos M et al (2014) Adverse weather conditions for European wheat production will become more frequent with climate change. Nat Clim Chang 4:637–643
Virkajärvi P, Pakarinen K, Hyrkäs M et al (2012) Tiller characteristics of timothy and tall fescue in relation to herbage mass accumulation. Crop Sci 52:970–980
Volenec JJ, Nelson CJ (2007) Physiology of forage plants. In: Barnes R, Nelson J, Moore K et al (eds) Forages—the science of grassland agriculture. Volume II. Blackwell Publishing Professional, Ames, IA, pp. 37–52
Wang J, Cui L, Wang Y et al (2009) Growth, lipid peroxidation and photosynthesis in two tall fescue cultivars differing in heat tolerance. Biol Plant 53:237–242
Wilman D, Gao Y, Leitch M (1998) Some differences between eight grasses within the Lolium-Festuca complex when grown in conditions of severe water shortage. Grass Forage Sci 53:57–65
Ylhäisi J, Tietäväinen H, Peltonen-Sainio P et al (2010) Growing season precipitation in Finland under recent and projected climate. Nat Hazards Earth Syst Sci 10:1563–1574
Ylimäki A (1967) Root rot as a cause of red clover decline in leys in Finland. Ann Agric Fenn 6:1–59
Acknowledgements
We thank the Academy of Finland for financing the projects ADIOSO (decision no. 255954) and the A-La-Carte (decision no. 140870). We are grateful to the providers of the data of the MTT Official Cultivar Trials and the Finnish Meteorological Institute.
Author information
Authors and Affiliations
Corresponding author
Compliance with ethical standards
Online Resource 1
(DOCX 169 kb)
Rights and permissions
Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.
About this article
Cite this article
Hanna, M., Janne, K., Perttu, V. et al. Gaps in the capacity of modern forage crops to adapt to the changing climate in northern Europe. Mitig Adapt Strateg Glob Change 23, 81–100 (2018). https://doi.org/10.1007/s11027-016-9729-5
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11027-016-9729-5