Journal of Agricultural Science (2006), 144, 381392. f 2006 Cambridge University Press doi:10.

1017/S0021859606006423 Printed in the United Kingdom

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CENTENARY REVIEW

A physiological analysis of oilseed rape yields : Past and future

P. M . B E R R Y 1* A N D J. H . S P I N K 2
1

ADAS, High Mowthorpe, Malton, North Yorkshire, UK 2 ADAS, Rosemaund, Hereford, Herefordshire, UK

(Revised MS received 10 August 2006; First published online 18 September 2006)

SUMMARY Oilseed rape yields on farms have not increased in several countries, including the UK, since the mid 1980s. This may be because the yield potential for the environment in these countries has been reached, or due to a lack of genetic improvement, or due to changes in the environment of the growing crop caused by crop management practices. The present paper investigates which of these factors may be causing the yield of farm crops in the UK to remain at 3 t/ha. The yield potential for the UK that would be possible by combining the best characteristics that have been observed with the best crop management is estimated to be signicantly greater than average farm yields at 6.5 t/ha (90 g/kg moisture content). In order to achieve 6.5 t/ha, a crop would have a ower cover of less than 0.4 to facilitate the production of 130 000 seeds/m2. Seed lling must last 46 days and have a solar radiation use eciency of 0.75 g of seed/MJ. A tenth of the yield must come from stem reserves and the seed weight must be 5.0 mg. All of these characteristics have been achieved ; therefore the challenge lies in combining these traits within the same crop. The ultimate yield potential for water retentive soils in the UK is estimated at 9.2 t/ha. This would require new characteristics to be bred into the crop and represents a long-term target. In the UK, new oilseed rape varieties are introduced each year and the yield of these varieties under optimum growing conditions is estimated to have increased by 62 kg/ha/year between 1978 and 2005. Lack of genetic improvement is therefore unlikely to explain the halt in farm yields. There have been trends for farm crops to be grown in shorter rotations, established using minimal cultivations rather than ploughing and to receive less nitrogen fertilizer. Sulphur applications have increased, but probably have not kept pace with the reduction in deposition from the atmosphere. Fungicide applications to farm crops are less than applied to the variety testing system and are unlikely to oer complete disease control. It is concluded that a combination of these crop management factors has caused the halt in yield improvement on UK farms.

INTRODUCTION Four species of brassica have been widely cultivated as oilseed crops : Brassica napus (Swede rape), B. rapa (Turnip rape), B. juncea (Indian mustard) and B. carinata (Ethiopian mustard). When grown for oilseed, these species are commonly known as rapeseed or oilseed rape. The genomic relationships between these brassica species are well understood (Downey & Rimmer 1993). Brassica rapa (n=10 ; A), B. nigra (n=8 ; B) and B. oleracea (n=9 ; C) are the
* To whom all correspondence should be addressed. Email: pete.berry@adas.co.uk

primary species. Brassica juncea (n=18 ; AB), B. napus (n=19 ; AC) and B. carinata (n=17 ; BC) are amphidiploids resulting from crosses between corresponding pairs of the primary species. The winter form of B. napus is the most common rapeseed grown in Europe and China. In cooler areas of northern Europe and in Canada, the summer form of B. napus or the winter or summer forms of B. rapa are grown. In Australia, where water is often limiting, the summer form of B. napus has become the most frequently grown rapeseed. Brassica juncea is commonly grown in India and parts of China. Brassica carinata is grown in North East Africa.

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Table 1. Area of rapeseed (1000 ha)

1965 World China India Canada Germany France Australia UK Poland 7065 1842 2910 581 165 173 0 1 274 1975 9911 2314 3680 1829 222 282 16 39 309 1985 14 756 4494 3987 2783 410 474 74 296 467 1995 23 816 6907 6060 5273 974 864 377 439 606 2005 26 951 7220 6800 5154 1345 1211 1080 603 544

Data source: FAOSTAT Data (2006). www.faostat.fao.org.

The global area of rapeseed has increased from just over 7 million ha in 1965 to almost 27 million ha in 2005 (Table 1). This increase represents a doubling of the area grown every 20 years. In 2005, China grew the greatest proportion of the worlds rapeseed area (0.27), followed by India (0.25), Canada (0.19), Europe (0.16) and Australia (0.04). Since 1995, the greatest proportional increases in area grown have taken place in Australia and Europe. In 2000, rapeseed provided 0.16 of the vegetable oil used for human and animal feed and industrial use (www. usda.gov). Oil from rapeseed was the third most important source of vegetable oil behind soybeans (0.31) and palm (0.28). Rapeseed also provides about 0.12 of the world supply of protein meal (www. usda.gov). YIELD TRENDS Yields vary widely around the world. Since 2000, the average yields have been 3 t/ha or more in the UK, Germany and France, 2.4 t/ha in Poland, and between 1.6 and 1.0 t/ha in China, Canada, Australia and India. The average world yield of rapeseed has increased from about 0.6 t/ha in the early 1960s to 1.6 t/ha in the 2000s. The average yield increase per year has been 26 kg/ha. This rate of increase has been fairly constant since the 1960s and shows no signs of levelling o (Fig. 1). Similarly consistent yield increases have been repeated in some countries including China, where the mean annual increase has been 30 kg/ha, India (15 kg/ha), Canada (14 kg/ha) and Germany (39 kg/ha). In France, Australia, the UK and Poland there appears to have been a reduction or halt in the rate of yield increase (Fig. 1). In these countries, the average annual yield increase up to 1985 was between 28 and 65 kg/ha (Table 2). However, no signicant yield trend can be detected for any of these countries after 1985 (Table 2). This pattern of annual yield for the UK is consistent with other sources of data (e.g. statistics from the

Department for Environment, Food and Rural Aaris, DEFRA ; http ://statistics.defra.gov.uk). It is clear, therefore, that farm yields have stopped increasing in several countries. This may be because the yield potential for the environments of these countries has been reached, making further yield increases impossible. Alternatively, the yield potential may be greater, but this is not being realized due to a lack of genetic progress. A nal possibility is that changes to the growing environment, as inuenced by agronomic practice or the climate, are preventing the genetic potential from being realized. The present review assesses which of these three scenarios is the most likely to be the cause of the apparent lack of yield improvement by using the UK as a case study. POTENTIAL YIELD Yield is dened as the weight of seed measured at 90 g/kg moisture content. Calculations of potential yields are by denition speculative. The present paper will calculate two potential yields : a conservative estimate based on characteristics already observed, but which have not necessarily been combined in the same crop, and an ultimate potential yield which relies on new traits being developed by plant breeders. Potential yield is dened here as the yield that could be produced by a combination of the best germplasm, combined with the best management, on deep soil (>1.5 m) and in an environment with the current average radiation, temperature and rainfall for the UK. The yield of oilseed rape may be usefully split into two main components : seed number/m2 and individual seed weight. This section investigates the potential for increasing both of these components. Seed number/m2 The number of seeds/m2 determines the size of the sink. This character has frequently been demonstrated

Oilseed rape yield review

(a) 4 3 Yield (t/ha) Yield (t/ha) 2 1 0 1960 1970 1980 1990 2000 (d ) 4 3 Yield (t/ha) Yield (t/ha) 2 1 0 1960 1970 1980 1990 2000 (g) 4 3 Yield (t/ha) Yield (t/ha) 2 1 0 1960 1970 1980 1990 2000 (h ) 4 3 2 1 0 1960 1970 1980 1990 2000 Yield (t/ha) (e) 4 3 2 1 0 1960 1970 1980 1990 2000 (i ) 4 3 2 1 Yield (t/ha) (b) 4 3 Yield (t/ha) 2 1 0 1960 1970 1980 1990 2000 (f) 4 3 2 1 (c) 4 3 2 1 0 1960 1970 1980 1990 2000

383

0 1960 1970 1980 1990 2000

0 1960 1970 1980 1990 2000

Fig. 1. Yield trends for (a) World, (b) China, (c) India, (d ) Canada, (e) Germany, ( f ) France, (g) Australia, (h) UK, (i ) Poland. Data Source: FAOSTAT Data (2006). All yields are calculated on a minimum area of 5000 ha.

to be the most important yield component. For example, it was shown to account for 0.85 of yield variation by Mendham et al. (1981). The number of seeds/m2 is determined during a critical phase for pod and seed abortion lasting about 300 xCd after mid-owering (Mendham et al. 1981; Leterme 1988). In most eld situations this equates to about 1925 days. Pod and seed survival have been shown to be related to the amount of radiation intercepted by

photosynthetic tissue per ower and per pod respectively during this critical period (Mendham et al. 1981; Leterme 1988). The radiation intercepted by green tissue at this time is severely reduced by the layer of owers, which absorb and reect radiation. Flower cover has been measured at 0.50 during mid owering across several varieties (Yates & Stevens 1987). Mendham et al. (1981) measured a similar sized ower cover of 0.60. The owers of a crop with a

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Table 2. Yield trends (kg/year)

19611985 World China India Canada Germany France Australia United Kingdom Poland +26 +34 +9 +16 +42 +40 +28 +65 +29 P <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.05 <0.001 <0.01 19862005 +22 +32 +13 +13 +32 P <0.001 <0.001 <0.01 <0.01 <0.05 >0.05 >0.05 >0.05 >0.05

30 25 Seeds per pod 20 15 10 5 0 0 5000 10000 pods/m2

100000 90000 80000 70000 50000 40000 30000 20000 10000 0 15000 seeds/m2 60000

Data source: FAOSTAT Data (2006). www.faostat.fao.org.

ower cover of 0.62 at mid-owering were measured to absorb and reect 0.58 of photosynthetically active variation (Yates & Stevens 1987). The components of seed number/m2, pod number/m2 and seed number/pod are negatively related (Fig. 2). This relationship results in an optimum fertile pod number of between 6000 and 8000 pods/m2 (Fig. 2). It is likely that the canopy size of crops with fewer than 6000 pods/m2 is too small to trap all incident radiation, which results in fewer seeds set/m2. For crops with more than 8000 pods/m2 it is likely that the thickness of the owering layer reduces the amount of radiation reaching the photosynthetic tissues, which reduces the number of seeds/pod and seeds/m2. In order to maximize seed number/m2, the amount of photo-assimilate produced during the period of seed determination must be maximized. Methods of achieving this include bringing owering forward, reducing the amount of light reected by the owering layer and increasing the leaf area. Bringing owering forward into cooler days will increase the number of days required to achieve the 300 xCd period over which seed number is determined which will increase the radiation received during this period. Anecdotal evidence indicates that yield losses from frost damage are rare in the UK and so there appears to be scope to bring owering forward by a modest amount of about one week without increasing the risk of frost damage to levels that will reduce yield. This may be done by choosing early developing varieties and/or early sowing. If the date of mid-owering of current crops is taken as 1 May then bringing owering forward by 7 days will increase the radiation received by the crop during the seed determination period by about 2 %. Reducing the amount of light reected by the owering layer without reducing the green area of the crop may be achieved by reducing the area of individual petals. Apetalous lines oer a potential solution and have been shown to increase yield in

Fig. 2. Seeds per pod plotted against pods/m2 (crosses) for sowing date, seed rate and N fertilizer treatments at two UK sites in 1996, 1997 and 1998 (Lunn et al. 2001). Best t line (dashed); y=13.2 Ln(x)+130.2, R2=0.87 (P<0.001). Relationship between seeds/m2 and pods/m2 (solid line) calculated from relationship between seeds/pod and pods/m2.

Australia (Rao et al. 1991). In the UK however, an apetalous variety did not increase yield (Fray et al. 1996). This may have been due to the production of more pods and branches rather than the anticipated increase in seeds/pod, or that the lines used were early breeding material that was not well adapted (P. Werner, CPB Twyfords, personal communication). More recent work has indicated that apetalous traits could be an advantage in the UK (Evans et al. 2003). Flower cover at mid owering has been shown to vary by 0.50 between varieties (Yates & Stevens 1987). This variation was due to a combination of a variation in petal size of more than 0.50 and variation in ower number. In Yates & Stevens (1987), the varieties with moderate ower covers of 0.380.50 had a greater seed yield than varieties with a ower cover of greater than 0.50. It was hypothesized that varieties with ower covers of below 0.38 did not follow this trend because they had insucient pods. Excessive ower cover can also be reduced through agronomic means, e.g. by avoiding very early sowing, using lower seed rates and applying plant growth regulators (Lunn et al. 2003). Reducing the ower cover from 0.50 of a typical canopy to 0.38 will increase the amount of radiation received by the green tissue by 25 %. During owering, crops with an optimum number of pods have about 2.5 units of leaf area and 1.5 units of stem area (Lunn et al. 2001). The radiation use eciency per unit area of leaf material is three times greater than stem tissue (Major 1977). If leaf area can be increased to 3 units by increasing the size and

Oilseed rape yield review duration of the leaves, and the stem area can be reduced to 1 unit through shortening, then the overall radiation use eciency of the leaf and stems canopy would increase by about 12 %. If owering can be brought forward by 1 week, ower cover reduced by 0.25 and leaf area increased, then the increase in photo-assimilate during the period of seed determination is estimated to increase by 39 %. Applying this to crops with an optimum number of pods would increase the number of seeds from 93 000 to 130 000/m2. Crops with 130 000 seeds/ m2 have been recorded in the UK (Fray et al. 1996) and in Australia (Rao et al. 1991). Each pod would contain 19 seeds, which is well below the maximum observed of 30 (Mendham & Salisbury 1995). Seed growth The period from mid-owering to physiological maturity has been shown to last for 715 xCd above 4.2 xC in cv. Victor grown in the UK (Mendham et al. 1981). If mid-owering occurs on 23 April, then physiological maturity will occur on the 1 July in average UK temperatures if the relationship for cv. Victor is used. The end of seed number determination has been estimated to occur on 18 May. This gives a seed growth period of 43 days. Results from the national variety testing system show that scores for the earliness to owering and earliness to maturity are not well correlated (Anon. 2006). Moderate-late maturing varieties can either be early or late owering. Some of this variation may be caused by variation in the duration of owering, but it seems likely that there must be some varietal dierences in the duration of seed lling. A seed-lling period of 46 days therefore seems realistic. The solar radiation use eciency (RUE) during seed lling has been measured at between 0.4 g/MJ (Habekotte 1997) to 0.75 g/MJ (Dreccer et al. 2000). This compares with RUEs before owering of 1.2 g/ MJ (Mendham et al. 1981), 1.35 g/MJ (Habekotte 1997; Justes et al. 2000) and 1.7 MJ/g (Rao et al. 1991). The pre-owering values are within the range of RUEs commonly observed for C3 crops within temperate environments (Sinclair & Muchow 1999). RUE is less during seed lling for two reasons : rstly, 45 % more assimilate is required to produce each gram of oil-rich seed compared with pre-anthesis biomass (Sinclair & de Wit 1975). Accounting for this means that the post-anthesis RUEs are equivalent to pre-anthesis RUEs of 0.58 to 1.09 g/MJ. Secondly, pods have a photosynthetic capacity that is estimated at between 0.50 and 0.70 of that of leaves (Major 1977; Gammelvind et al. 1996). Therefore, it is clear that the RUE during seed lling cannot be expected to match that attained before owering. A realistic expectation of the RUE that could be achieved during seed lling is given below.

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The photosynthetic capacity of stems and pods has been estimated at 0.37 and 0.67, respectively, of the leaves (Major 1977). Therefore, maximizing the proportion of leaves within the canopy will increase overall photosynthesis. At owering, the optimum canopy structure has been estimated to have a green area index of about four (Lunn et al. 2003) of which three units are leaf and one unit is stem. During owering and seed determination, the leaf area decreases and the pod area increases resulting in little change in the overall GAI. In less dense canopies at the beginning of seed lling, leaves can make up 0.30 of the total green area and can persist throughout most of the seed lling period (McWilliam et al. 1995; Staord 1996). The green area of the stems and pods then change little during seed lling (Norton et al. 1991). It seems reasonable to assume that the GAI of an optimum canopy at the start of seed ll is four units and the leaf area will decline linearly from 0.30 to zero at the end of seed lling. Stem and pod areas will remain constant at one unit and 1.8 units respectively. Averaged over the whole seed lling period, leaves would represent 0.18 of the total green area, stems 0.29 and pods 0.53. This compares with pre-owering when leaf area makes up 0.75 of the area and stems 0.25. If it is assumed that the amount of light intercepted is proportional to the area of each tissue component and photosynthetic capacity is as estimated by Major (1977), then the RUE during seed lling is estimated to be 0.76 of the RUE before owering. Accounting for the extra energy costs of forming oil-rich seed, reduces this to 0.53. The maximum pre-owering RUE has been measured at 1.7 g/MJ (Rao et al. 1991), so it seems reasonable to assume that the post-owering RUE measured by Dreccer et al. (2000) of 0.75 g/MJ is a realistic target. Attaining this RUE target could be further facilitated by using erectophile pods (Fray et al. 1996) and by avoiding lodging. Lodged crops have been shown to reduce yield by between 16 and 50 % (Baylis & Wright 1990; Armstrong & Nichol 1991). Lodging compresses the oilseed rape canopy and forces the photosynthetic tissues to adopt a horizontal posture. This reduces the eciency with which the crop is able to use the available light because the upper pods easily become light saturated due to their modest photosynthetic capacity and the lower pods experience low levels of light. In oilseed rape, lodging can occur by either anchorage failure or stem buckling/ breakage (Goodman et al. 2001). Lodging risk can be eectively reduced with shorter crops, stier stems and stier, longer tap roots (Goodman et al. 2001). Breeders could exploit semi-dwarfs to reduce lodging risk. There may also be genetic variation in stem stiness and root properties as found in wheat by Berry et al. (2003).

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The proportion of incident radiation intercepted at the beginning of seed lling by a crop with a GAI of 4 and extinction coecient of 0.6 is 0.91. This is expected to decrease to 0.81 at the end of seed lling, due to senescence of the leaf canopy. This results in an average proportion of light intercepted during grain lling of 0.86. The average amount of incident radiation in the UK during the proposed 46-day seed lling period has been calculated at 868 MJ/m2. If 0.86 of the incident radiation is intercepted by the crop and if a RUE of 0.75 g/MJ can be achieved, then the amount of dry matter accumulated (allowing for the oil-rich nature of the seed) during the seed lling period will be 560 g/m2. At the start of seed lling the pods have reached their nal length, but pods still accumulate some biomass during seed lling. It has been estimated that pod wall growth accounts for between 0.05 and 0.25 of the biomass 1993; accumulated during seed lling (Habekotte Hocking & Mason 1993). Assuming the lower amount gives a seed biomass of 532 g/m2. Water-soluble carbohydrate accumulated before owering and predominantly stored in the stem has been estimated to contribute a negligible amount (Staord 1996), 0.10 (Mendham 1995) or 0.12 1993) to nal yield. If stem reserves con(Habekotte tribute 0.10 of nal yield then, due to the increased energy content of the seed, 85 g/m2 of stem reserves must be relocated to increase seed yield by 59 g/m2 to 591 g/m2. Stem reserves have been measured at 1993) so this between 40 and 110 g/m2 (Habekotte amount of relocation seems feasible. A crop yielding 591 g/m2 with 130 000 seeds/m2 would have a mean seed dry weight of 4.55 mg, which is well within the range of observed seed weights (Mendham & Salisbury 1995). At 910 g DM/kg the theoretical potential yield of oilseed rape in the UK is 6.49 t/ha. The amount of water required to produce a crop with a dry matter yield of 591 g/m2 can be estimated by assuming the commonly observed water use eciency of 5 g of carbohydrate dry matter/m2/litre of water (Green et al. 1983). Taking into account the higher energy demand for the production of oil, it may be assumed that this will decrease to 3.4 g/m2/l for the seed (Penning et al. 1989). If it is assumed that the stem and leaf biomass of a high-yielding crop remains the same as a current average crop at 700 g/m2 and the pod biomass is 0.50 of the seed 1993), then the total amount of weight (Habekotte non-seed biomass will be 996 g/m2. This crop will require 355 mm of rainfall or soil water storage, after accounting for the contribution of stem reserves. The majority of this water will be required between the onset of stem extension in mid March and canopy senescence on the 1 July. If it is assumed that the crop biomass in mid March is 200 g/m2, then the crop will require 315 mm during the following 3.5 months.

Assuming that the soil is at eld capacity in mid March, then the available water capacity of the soil to a depth of 1.5 m would be between 210 and 330 mm, depending on soil type. Using information about the root length density of oilseed rape (Barraclough 1989) together with relationships linking root length density with water extraction for cereals (King et al. 2004), it is estimated that oilseed rape roots may extract about 0.68 of available water to a depth of 1.5 m. This means that oilseed rape crops could extract between 143 and 224 mm of water from the soil. This leaves a shortfall of between 91 and 172 mm that must be supplied by rainfall between mid March and 2 July. Average monthly rainfall varies between 45 mm in the east of the UK and 83 mm in the south and west. Therefore, in a year with average rainfall, the water supply would prevent the maximum yield from being attained in crops grown on low moisture retentive soils such as sandy loams in the east where the maximum yield would be 6.0 t/ha (at 910 g DM/kg). At this point it must be recognized that oilseed rape is mainly grown on heavy soils. This conservative estimate of yield potential of 6.5 t/ha should be achievable given that all of the characteristics have been reported in peer-reviewed literature or are realistic for current varieties. The challenge lies with the breeders to combine these traits in the same crop and with agronomists to ensure that crop management is optimal for achieving the characteristics. A comparison of the characteristics of a crop yielding 6.5 t/ha with characteristics of a crop yielding the UK farm average yield of 3 t/ha illustrates that several traits must undergo a signicant amount of improvement (Table 3). However, farm yields above 5 t/ha have been observed and this suggests that parts of farms/elds could be approaching 6 t/ha. Therefore, the estimated potential yield with current germplasm of 6.5 t/ha does not seem unrealistic. In order to estimate the ultimate yield potential, it must be assumed that plant breeders can improve some plant characteristics outside the range that has been observed to date. Clearly, the estimate of potential yield presented in the current paper is much more speculative. Key characteristics for which improvement would have a large eect on yield include the seed number/m2, duration of seed lling, radiation use eciency during seed lling and the proportion of pre-owering assimilate transferred to the seeds. Apetalous varieties should provide a route for reducing the amount of light reected by the ower canopy, which should increase the number of seeds set to at least 150 000/m2. Genetic variation in the duration of seed lling appears to exist and further selection could increase this period to 50 days. The low radiation use eciency of stems and pods has been linked with the low density of stomata of these tissues (Major 1977). If this could be increased to the level

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Table 3. Characteristics of crops yielding the UK farm average, a potential yield for existing germplasm and a potential yield with breeding advances
Yield potential for existing germplasm 23 Apr 0.38 130 000 18 May 46 days 0.86 0.05 0.10 0.75 g/MJ 5.0 mg 6.5 t/ha 316 mm

Typical farm crop Date of mid-owering Proportion ower cover Number of seeds/m2 Seed lling traits Start date Duration Light intercepted by green tissue Assimilate used for pod growth Contribution of stem reserves RUE during seed lling Individual seed weight (910 g/kg dry matter) *Yield (910 g/kg dry matter) Water use from start of stem extension 1 May 0.60 80 000 23 May 40 days 0.86 0.15 0.05 0.47 g/MJ 3.8 mg 3.0 t/ha 203 mm

Ultimate yield potential 23 Apr 0.30 150 000 18 May 50 days 0.86 0.05 0.20 0.88 g/MJ 6.1 mg 9.2 t/ha 393 mm

* The same oil content has been assumed for both yield estimates.

of leaves then RUE during seed lling would increase from 0.75 to 1 g/MJ. This is a large step increase and it may be more reasonable to set a more modest target of 0.88 g/MJ. Relatively little is known about the amount of pre-owering assimilate that is used for seed growth, with estimates ranging from 0 to 0.10 of yield coming from pre-owering assimilate. These rates are signicantly less than wheat for which preowering assimilate can account for as much as 0.30 of yield. Therefore, a future target of 0.15 is set for oilseed rape. Using these characteristics with the calculations described previously gives an ultimate yield potential of 9.2 t/ha (Table 3). However, to achieve this yield level would require 393 mm of water stored in soil and rain after the start of stem extension. Water availability will therefore limit yield in many areas, with yield potential determined by the water-holding capacity of the soil and rainfall. In the east, where rainfall is low, this yield could only be achieved on loam, silt loam, silt clay loam and clay loam soils. The yield potential decreases to 8.65 t/ha on sandy clay loams, 7.4 t/ha on clays and silt clays and to 6.73 t/ha on sandy loams. Thus, the importance of improving root length density at depth so that more than 0.68 of available water can be extracted from the top 1.5 m of soil is apparent. These estimates of potential yield illustrate that the yields achieved on farm are signicantly below the yield potential in the UK environment. The lack of yield improvement must therefore be due to either a lack of genetic improvement or restrictions imposed by the growing environment through crop management. These issues are discussed in the next sections.

GENETIC CONSTRAINTS TO YIELD IMPROVEMENT Since the late 1970s, the UK has employed a system of evaluating the performance of current and new oilseed rape varieties (e.g. Anon. 2006). Each year between three and 23 varieties have been grown within replicated trials at about 20 sites in oilseed rape growing areas within the UK. The crop agronomy used is designed to maximize the chance that the genetic potential for each variety is realized. Thus, the availability of nutrients is not limiting and weeds, diseases and pests are generally fully controlled through the use of cultural practices and prophylactic use of chemical controls. Surveys have shown that uptake of new oilseed rape varieties by farmers is rapid (Sylvester-Bradley et al. 2002), so the varieties used to calculate the yields in this testing system will approximate to the same varieties grown on farms 1 or 2 years later. The mean yield of the varieties grown in the testing system in each year increased by an average of 62 kg/ha between 1978 and 2005 (Fig. 3). The trend for increased yield has been consistent apart from low yielding years in 2000, 2001 and 2003. The highest average yield of any year was in 2005, which indicates that yields are continuing to increase. The upward yield trend is impressive when it is considered that the breeding programmes suered major interruptions to breed for low erucic acid in the 1970s and low glucosinolate in the mid 1980s. The development of low glucosinolate lines initially caused a reduction in oil concentration to below 400 g/kg, on average. However, between 1986 and 1990 the average oil

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quick to adopt new varieties it therefore appears that the lack of yield increases on-farm since 1985 has not been due to a lack of germplasm improvement. CROP MANAGEMENT CONSTRAINTS TO YIELD IMPROVEMENT

Yield (t/ha)

0 1956

1966

1976

1986

1996

2006

Fig. 3. Average yield of varieties evaluated in the national variety testing system (squares) (Anon. 2006) and average farm yield (crosses) (FAOSTAT Data, 2006).

concentration climbed to 430 g/kg and has remained at 430440 g/kg since then. Thus, it is clear that the increase in seed yield has not been achieved at the expense of oil concentration. The rate of yield increase within the testing system is very similar to the rate of yield increase of farm yields between 1967 and 1985 (Table 2 ; Fig. 3). Since 1985, the farm yields appear to have been static whereas the yields of the testing system clearly continue to increase (Fig. 3). The UK farm yields include spring-sown crops, which are inherently lower yielding. The proportion of spring crops has shrunk in recent years from 0.10 to less than 0.05. Therefore, the dierence in yield trends cannot be explained by an increase in the proportion of spring-sown crops. Industrial oilseed rape is grown on set-aside land and accounts for 0.100.20 of the oilseed rape crop. It has been suggested that industrial crops may be managed less intensively than oilseed rape grown for human consumption. However, this also does not appear to account for the dierence as industrial crop yields have averaged just 0.2 t/ha less than conventional crops over the last 7 years data (http://statistics. defra.gov.uk). Trials carried out without fungicides within the variety testing system have shown that disease resistance of some varieties can break down over several years (Spink & Berry 2005). This could help explain why farm yields have not improved. However, with a full programme of fungicides very few varieties grown in the national variety testing system show any slippage in yield over time. This indicates that farmers should be able to counter any breakdown in genetic resistance through increased fungicide use. Given that UK farmers have been

The previous analyses have demonstrated that farm yields are not limited by the yield potential for the UK environment and that the genetic potential has been consistently improving. Therefore, changes in the management of farm crops, particularly those that result in dierent management to that used in the national variety testing system, are most likely to explain the lack of yield increase in farm crops (Fig. 3). Identifying these dierences should help explain why farm yields have stopped increasing. Several management changes can be identied including rotation length, cultivations, fungicide use, fertilizer use and sowing date. The frequency with which oilseed rape is grown in arable rotations has increased because it is commonly the second most protable combinable crop after rst winter wheat and is an eective break for wheat. The frequency with which oilseed rape is grown one year in three had increased from 0.08 in 1990 to 0.15 in 1994 (Turner & Hardwick 1995) and then to 0.23 in 2003 (J. Turner, personal communication). Shorter rotations will increase the likelihood of diseases and disease-associated yield loss. Sieling & Christen (1997) showed that oilseed rape grown every 2 years yielded about 0.5 t/ha less than when it was grown once every 3 years. Years with low farm yields have been correlated with years when the incidence of phoma canker (Leptosphaeria maculans (Phoma lingam)), light leaf spot (Pyrenopeziza brassicae) and dark pod spot (Altenaria brassicae) were high (Booth et al. 2005). Between 1987 and 1994, less than half of commercial crops received a fungicide in autumn or spring to control light leaf spot or phoma and a quarter to half received a fungicide at or post owering to control stem rot (Sclerotinia scleretiorum) and dark pod spot (Fitt et al. 1997). Since 1998, the number of crops receiving a fungicide has stabilized at 0.90 (Turner et al. 2002). Disease induced yield losses increased steadily between 1997 and 2000 despite the use of fungicides (Turner et al. 2002) and it is clear that dose and timing of the fungicide is not always optimal (Fitt et al. 1997; Turner et al. 2002). The increase in disease may be attributed to warmer weather or the breakdown of varietal resistance. It is probable that yields under the testing system will be aected less by diseases due to a more robust programme of fungicides compared with use on commercial crops. This includes three compulsory fungicides in the autumn, beginning of stem extension and at owering, with a further three fungicide

Oilseed rape yield review applications possible depending on disease pressure. This contrasts with an average of just 1.8 fungicide sprays applied to farm crops in 2004 (Garthwaite et al. 2004). Many farm crops are now autocast directly into the preceding cereal crop, direct drilled or established after minimum tillage as compared with ploughing which has traditionally been the main method of cultivation on farms and is used in the variety testing trials. Direct drilling or autocasting have been shown to result in lower yields compared with ploughing, but yields from crops established using minimal cultivations are similar to those established with ploughing (Sauzet et al. 2003 ; Bowerman et al. 1995). Conversely, Christensen et al. (2003) found no detrimental eect of direct drilling on yield. It appears direct drilling and autocasting are less eective when there are large amounts of poorly chopped straw from the previous crop, wet conditions or wet and poor soil structure. Non-burial of the residues of the previous crop has also been shown to increase lodging and phoma infection (Sauzet et al. 2003) and increase slug damage, which may cause suboptimal plant populations to be established. Many farm crops are established 23 weeks earlier than crops grown in the national variety testing system. Early sowing has commonly been shown to reduce seed yields (Mendham et al. 1981; Jenkins & Leitch 1986; Leach et al. 1994, 1999). Early sowing also increases the risk of pests such as cabbage root y (Delia radicum L.), light leaf spot and aphids and viruses. Extensive trials work in Germany showed that drilling 2 weeks earlier (late August compared with early September) reduced yields by 0.1 to 0.2 t/ha (Baer & Frauen 2003). This is thought to be due to the production of an over-thick canopy from early drilled crops which reduces the number of seeds set. Oilseed rape crops yielding 3 t/ha require about 50 kg of sulphur/ha (McGrath & Zhao 1996). In the past, atmospheric depositions have largely met this requirement, but between 1970 and 2002 SO2 emissions have decreased by 82 % (source : NET CEN). The proportion of farm crops that had sulphur applied increased from 0.10 to 0.30 between 1993 and 1996, then remained constant at 0.30 until 2001. Since 2001, the proportion of crops receiving sulphur has risen sharply to 0.50. The average application rate has risen steadily from less than 15 kg S/ha in 1993 to 30 kg S/ha in 2003. Many oilseed rape crops are responsive to S fertilizer, with yield increases of 0.7 to 1.6 t/ha recorded in response to 40 kg S/ha on a sandy soil (McGrath & Zhao 1996). An extra application of sulphur began to be applied to the testing system trials during the early 1990s (S. Kightley, personal communication) and a minimum of 30 kg S/ha is currently recommended. Farmers have therefore been slow to make up for the reduction in sulphur
Average application rate (kg ha1) 300 250 200 150 100 50 0 1973 1978 1983 1988 1993 1998

389

2003

Fig. 4. Applications of nitrogen (solid line), phosphate P2O5 (dashed line) and potassium (K2O) (dotted line) to oilseed rape in the UK. Data source: Goodlass et al. (2003).

emissions and many probably still do not apply enough to prevent sulphur deciency from limiting yield potential. Nitrogen fertilizer applications to commercial oilseed rape crops increased from 250 kg N/ha to 280 kg N/ha between 1980 and 1984 then decreased steadily to 180 kg N/ha by 1993 (Fig. 4). Since then, application rates have risen slowly to current levels of about 200 kg N/ha (Goodlass et al. 2003). The reduction in N fertilizer use during the late 1980s and early 90s was largely a result of falling oilseed prices. Nitrogen fertilizer applications to the testing system trials include 30 kg N/ha in the autumn and then follow the farm practice on which the trial is situated. Thus, it is likely that testing system trials have also undergone a reduction in N applications since the mid 1980s. Applications of potassium and phosphate have also decreased slightly since the 1980s (Fig. 4). Farm oilseed rape crops now tend to be grown in shorter rotations, to be established using minimal cultivations or autocasted rather than after ploughing, and to receive less nitrogen fertilizer than they did 20 years ago. Sulphur applications have increased, but it is unlikely that they have kept pace with the reduction in depositions from the atmosphere over the same period. Fungicide applications to farm crops are signicantly less than applied to the variety testing system and are unlikely to oer sucient disease control when disease pressure is high. A signicant proportion of crops are sown earlier than trials in the national variety testing system and are likely to have a lower yield potential due to the production of a super-optimal canopy size. It seems likely that a combination of these factors has caused the lack of yield improvement on-farm since the mid 1980s. FUTURE PROGRESS Plant breeders have consistently increased the yield potential o new varieties towards the theoretical

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maximum of 9 t/ha since the 1970s. The most likely reason for the static farm yields since the mid 1980s is the employment of crop management practices that do not allow the genetic potential to be realized. Since the 1970s the real value of commodity crops, after accounting for ination, has declined steadily (Morris et al. 2005). Over the same period the cost of many of the inputs required for crop production have increased. This situation has reduced the economic level of inputs as illustrated by the uptake of autocasting and minimal tillage techniques and the decline in the rate of nitrogen fertilizer use since the mid 1980s. Further work must investigate whether similar factors are responsible for the lack of yield improvement in other countries including France, Poland and Australia. It is possible that the increase in genetic yield potential created by UK breeders has countered the eect of reducing inputs and has stopped farm yields from declining. Plant breeders may be able to increase farm yields by producing varieties that are able to perform well with fewer inputs, e.g. by increasing nitrogen use eciency, durable disease resistance and lodging resistance. However, these traits are probably under complex genetic control and will require signicant eort to improve them. If the price of oilseeds continue to decline in real terms then it

is likely that UK farm yields will remain static. However, if crop prices increase due to increasing demand for oilseeds for both food and fuel, this is likely to stimulate more attention to crop management inputs. This could trigger a rapid increase in yields as farmers close the large gap between farm yields and the genetic potential. Looking further ahead, climate change may have a signicant eect on yield trends in the UK. The Hadcm3 climate change model (http://www.metoce.gov.uk/research/hadleycentre/index.html) predicts that by 20702100 the mean temperatures in the UK will increase by 23 xC during March to May and by 25 xC during June to August. The amount of soil moisture available for plant uptake is not predicted to change for the March to May period and to decrease by between 0.5 mm and 2 mm per day for the June to August period. Assuming that the seed lling period lasts 715 xCd above 4.2 xC, then an increase in temperature of 3 xC would shorten the seed lling period by 13 days and reduce the amount of radiation intercepted by the crop during seed lling by 30 %. Carbon dioxide concentration in the atmosphere is predicted to increase over the same period, which is likely to increase RUE. It is therefore very dicult to predict how climate change may aect yield potential.

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