Nitrogen dynamics and nitrogen use efficiency of spring cereals under Finnish growing conditions

Nitrogen (N) is one of the main inputs in cereal cultivation and as more than half of the arable land in Finland is used for cereal production, N has contributed substantially to agricultural pollution through fertilizer leaching and runoff. Based on this global phenomenon, the European Community has launched several directives to reduce agricultural emissions to the environment. Trough such measures, and by using economic incentives, it is expected that northern European agricultural practices will, in the future, include reduced N fertilizer application rates. Reduced use of N fertilizer is likely to decrease both production costs and pollution, but could also result in reduced yields and quality if crops experience temporary N deficiency. Therefore, more efficient N use in cereal production, to minimize pollution risks and maximize farmer income, represents a current challenge for agronomic research in the northern growing areas. The main objective of this study was to determine the differences in nitrogen use efficiency (NUE) among spring cereals grown in Finland. Additional aims were to characterize the multiple roles of NUE by analysing the extent of variation in NUE and its component traits among different cultivars, and to understand how other physiological traits, especially radiation use efficiency (RUE) and light interception, affect and interact with the main components of NUE and contribute to differences among cultivars. This study included cultivars of barley (Hordeum vulgare L.), oat (Avena sativa L.) and wheat (Triticum aestivum L.). Field experiments were conducted between 2001 and 2004 at Jokioinen, in Finland. To determine differences in NUE among cultivars and gauge the achievements of plant breeding in NUE, 17-18 cultivars of each of the three cereal species released between 1909 and 2002 were studied. Responses to nitrogen of landraces, old cultivars and modern cultivars of each cereal species were evaluated under two N regimes (0 and 90 kg N ha-1). Results of the study revealed that modern wheat, oat and barley cultivars had similar NUE values under Finnish growing conditions and only results from a wider range of cultivars indicated that wheat cultivars could have lower NUE than the other species. There was a clear relationship between nitrogen uptake efficiency (UPE) and NUE in all species whereas nitrogen utilization efficiency (UTE) had a strong positive relationship with NUE only for oat. UTE was clearly lower in wheat than in other species. Other traits related to N translocation indicated that wheat also had a lower harvest index, nitrogen harvest index and nitrogen remobilisation efficiency and therefore its N translocation efficiency was confirmed to be very low. On the basis of these results there appears to be potential and also a need for improvement in NUE. These results may help understand the underlying physiological differences in NUE and could help to identify alternative production options, such as the different roles that species can play in crop rotations designed to meet the demands of modern agricultural practices.

Northern challanges for profitable production of quality naked oat

Naked oat (Avena sativa f.sp. nuda L.) is the highest quality cereal in northern growing conditions. However the cultivation area of naked oat is remarkably small. Major challenges for naked oat production are to observe its nakedness. The caryopsis of naked oat is sensitive to mechanical damage at harvest, especially at high grain moisture content. The greater the grain moisture content of naked oat at harvest, the more loses of germination capacity was caused by threshing. For producing high quality naked oat seed, it is recommended that harvesting be done at as low grain moisture content as possible. However, if this is not possible, better germination can be ensure with gentle harvest by reducing the cylinder speed. In spite of conventional oat s excellent fat and amino acid composition in animal feed use, as far as nutritional value is concerned, the total energy yield of oat is weaker than other cereals because of the hulls. Also with naked oat the dehulling is not complete, while hull content on different cultivars mostly varied between one to six percent. In addition to genotype, environmental conditions markedly control the expression of nakedness. Thresher settings had only limited effects on hull content. The function of hulls is to protect the groat, but this was confirmed only for Finnish, small grain, cultivar Lisbeth. The oat kernel is generally covered with fine silky hairs termed trichomes. The trichomes of naked oat are partly lost during threshing and handling of grains. Trichomes can cause itchiness in those handling the grains and also accumulate and form fine dust and can block-up machinery. The cultivars differed considerably in pubescence. Some thresher settings, including increased cylinder speed, slightly increased grain polishing such that grains had some areas completely free of trichomes. Adjusting thresher settings was generally not an efficient means of solving the problems associated with naked oat trichomes. The main differences in cultivation costs between naked and conventional oat lie in the amount of seeds required and the drying costs. The main differences affecting the economic result lie in market prices, yield level and feed value. The results indicate that naked oat is financially more profitable than conventional oat, when the crop is sold at a specific price at all yield levels and when the crop is used as feed at highest yield level. At lower yield levels, conventional oat is, in spite of its lower feed value, the more profitable option for feed use. Dehulled oat did not achieve the same economic result as naked oat, as the cost of dehulling, including the hull waste, was considerable. According to this study naked oat can be cultivated successfully under northern conditions, when taking into consideration the soft, naked grain through cultivation chain.

Undersowing in a northern climate: effects on spring cereal yield and risk of nitrate leaching

Disadvantages of invariable cereal cropping, concern of nutrient leaching and prices of nitrogen (N) fertilizer have all increased during last decades. An undersown crop, which grows together with a main crop and after harvest, could mitigate all those questions. The aim of this study was to develop undersowing in Finnish conditions, so that it suits for spring cereal farming as well as possible and enhances taking care of soil and environment, especially when control of N is concerned. In total, 17 plant species were undersown in spring cereals during the field experiments between 1991-1999 at four sites in South and Central Finland, but after selection, eight of them were studied more thoroughly. Two legumes, one grass species and one mixture of them were included in long-term trials in order to study annually repeated undersowing. Further, simultaneous broadcasting of seeds instead of separate undersowing was studied. Grain yield response and the capacity of the undersown crop to absorb soil N or fix N from atmosphere, and the release of N were of greatest interest. Seeding rates of undersown crops and N fertilization rates during annually repeated undersowing were also studied. Italian ryegrass (Lolium multiflorum Lam., IR) absorbed soil nitrate N (NO3-N) most efficiently in autumn and timothy (Phleum pratense L.) in spring. The capacity of other grass species to absorb N was low, or it was insufficient considering the negative effect on grain yield. Red clover (Trifolium pratense L.) and white clover (Trifolium repens L.) suited well in annually repeated undersowing, supplying fixed N for cereals without markedly increased risk of N leaching. Autumn oriented growth rhythm of the studied legumes was optimal for undersowing, whereas the growth rhythm of grasses was less suited but varied between species. A model of adaptive undersowing system was outlined in order to emphasize allocation of measures according needs. After defining the goal of undersowing, many decisions are to be done. When diminishing N leaching is primarily sought, a mixture of IR and timothy is advantageous. Clovers suit for replacing N fertilization, as the positive residual effect is greater than the negative effect caused by competition. A mixture of legume and non legume is a good choice when increased diversity is the main target. Seeding rate is an efficient means for adjusting competition and N effects. Broadcasting with soil covering equipment can be used to establish an undersown crop. In addition, timing and method of cover crop termination have an important role in the outcome. Continuous observing of the system is needed as for instance conditions significantly affect growth of undersown crop and on the other hand N release from crop residues may increase in long run.

Digestion capasity, nutrient digestibilities and physico-chemical conditions in the intestine influenced by the age of growing turkeys

Six experiments have been conducted to examine digestibility and feeding value of domestic Finnish fibre-rich cereals (barley and oats as compared to maize and wheat) and protein sources (rapeseed meal and cake, peas, faba beans, lupin seeds) for growing turkeys and to investigate effects of age of the birds (from 3 to 12 weeks of age) on digestion process and estimated nutrient digestibility and energy values. Besides, an objective of the study was to test applications of digestibility research methodology for turkeys. Total tract digestibility and apparent metabolizable energy (AME) was assayed in experimental cages using excreta collection, and a slaughter method was applied to sample small intestinal digesta for determination of apparent ileal crude protein digestibility (AICPD), jejuno-duodenal digesta viscosity and caecal volatile fatty acid (VFA) concentration. Digesta viscosity decreased and caecal VFA production increased with age of growing turkeys. Digesta retention times in the small intestine were generally longer in the older birds than in the younger ones. Crude fat digestibility and AME increased with age of growing turkeys, especially with viscous diets. AICPD seemed to decrease with age in most cases. Supplementation with β-gucanase-xylanase decreased viscosity, improved crude fat digestibility and metabolizable energy value and increased VFA production especially in barley-fed turkeys and especially in the young birds. Poor protein digestibility and low energy value of rapeseed meal and rapeseed cake decreased their feeding value for turkeys. In addition, a typical goitrogenic effect of rapeseed feeding was detected. Use of legume seeds as feed for growing turkeys is limited mostly by the low energy value in lupin seeds and the low ileal protein and amino acid digestibility in faba beans. Digestibility of fibre-rich protein sources was not improved with age of the turkeys. Euthanizing the turkeys for AICPD determination by carbon dioxide and bleeding led to lower digestibility values than mechanical stunning and cervical dislocation, suggesting inferiority of carbon dioxide stunning in experimental use. Comparison of AICPD and AME results obtained using different markers showed that considerable differences may occur, especially on total tract level, when acid-insoluble ash gave considerably lower AME values than titanium dioxide and chromic oxide.

Folates in rye: Determination and enhancement by food processing

The average daily intake of folate, one of the B vitamins, falls below recommendations among the Finnish population. Bread and cereals are the main sources of folate, rye being the most significant single source. Processing is a prerequisite for the consumption of whole grain rye; however, little is known about the effect of processing on folates. Moreover, data on the bioavailability of endogenous cereal folates are scarce. The aim of this study was to examine the variation in as well as the effect of fermentation, germination, and thermal processes on folate contents in rye. Bioavailability of endogenous rye folates was investigated in a four-week human intervention study. One of the objectives throughout the work was to optimise and evaluate analytical methods for determining folate contents in cereals. Affinity chromatographic purification followed by high-performance liquid chromatography (HPLC) was a suitable method for analysing cereal products for folate vitamers, and microbiological assay with Lactobacillus rhamnosus reliably quantified the total folate. However, HPLC gave approximately 30% lower results than the microbiological assay. The folate content of rye was high and could be further increased by targeted processing. The vitamer distribution of whole grain rye was characterised by a large proportion of formylated vitamers followed by 5-methyltetrahydrofolate. In sourdough fermentation of rye, the studied yeasts synthesized and lactic acid bacteria mainly depleted folate. Two endogenous bacteria isolated from rye flour were found to produce folate during fermentation. Inclusion of baker s yeast in sourdough fermentation raised the folate level so that the bread could contain more folate than the flour it was made of. Germination markedly increased the folate content of rye, with particularly high folate concentrations in hypocotylar roots. Thermal treatments caused significant folate losses but the preceding germination compensated well for the losses. In the bioavailability study, moderate amounts of endogenous folates in the form of different rye products and orange juice incorporated in the diet improved the folate status among healthy adults. Endogenous folates from rye and orange juice showed similar bioavailability to folic acid from fortified white bread. In brief, it was shown that the folate content of rye can be enhanced manifold by optimising and combining food processing techniques. This offers some practical means to increase the daily intake of folate in a bioavailable form.

Prolamin degradation in sourdoughs

This thesis examines protein behaviours that occur during cereal fermentations. The focus is on the prolamin degradation in sourdoughs. The thesis also looks at what happens to the oat globulins during an oat bran acidification process. The cereal prolamins are unique proteins in many respects. The wheat prolamins (glutenins and gliadins) are responsible for the formation of the gluten that provides the viscoelastic properties to wheat doughs whereas the rye prolamins (secalins) are unable to develop gluten-like structures. In addition, many baking technological features, such as flavour, shelf-life and dough properties are affected by the protein degradation that might occur during processing. On the other hand, the prolamins contain protein structures that are harmful to gluten sensitive people. It is thus evident that the degradation of the prolamins in sourdough processes may be approached from various aspects. This thesis describes some of these approaches. Four different cereal fermentations were carried out. Wheat sourdough (WSD) and rye sourdough (RSD) fermentations represented traditional sourdoughs. A germinated-wheat sourdough (GWSD) was a novel sourdough type that was prepared using germinated wheat grains that had high and diverse proteolytic activities. The oat bran fermentation (OBF) represented a fermentation system that lacked functional cereal proteases. The high molecular weight glutenins and rye secalins were degraded during the WSD and RSD fermentations, respectively. It was noteworthy that in WSD only a very limited degradation of the gliadins occurred. The gliadins were, however, hydrolysed very extensively during the GWSD fermentation. No protein degradation was observable in the OBF system. Instead the acidification altered the solubility of the oat globulins and this finally led to their aggregation. This thesis confirms that the endogenous proteases of cereals hydrolyse cereal prolamins in sourdoughs. The thesis also shows that the proteolytic activity of the used cereal raw material determines the extent of proteolysis that occurs in sourdough. This means that bakers may adjust the protein degradation in their sourdoughs by selecting the raw material based on its proteolytic activity. The thesis also demonstrates that by using germinated grains, with high and diverse proteolytic activity in sourdough preparations, the prolamins can be extensively degraded. Whether such highly proteolytic food technology could be used to manufacture new gluten-free cereal-based products for gluten sensitive people remains to be solved.

Bacillus cereus spores and cereulide in food-borne illness

B. cereus is a gram-positive bacterium that possesses two different forms of life:the large, rod-shaped cells (ca. 0.002 mm by 0.004 mm) that are able to propagate and the small (0.001 mm), oval shaped spores. The spores can survive in almost any environment for up to centuries without nourishment or water. They are insensitive towards most agents that normally kill bacteria: heating up to several hours at 90 ºC, radiation, disinfectants and extreme alkaline (≥ pH 13) and acid (≤ pH 1) environment. The spores are highly hydrophobic and therefore make them tend to stick to all kinds of surfaces, steel, plastics and live cells. In favorable conditions the spores of B. cereus may germinate into vegetative cells capable of producing food poisoning toxins. The toxins can be heat-labile protein formed after ingestion of the contaminated food, inside the gastrointestinal tract (diarrhoeal toxins), or heat stable peptides formed in the food (emesis causing toxin, cereulide). Cereulide cannot be inactivated in foods by cooking or any other procedure applicable on food. Cereulide in consumed food causes serious illness in human, even fatalities. In this thesis, B. cereus strains originating from different kinds of foods and environments and 8 different countries were inspected for their capability of forming cereulide. Of the 1041 isolates from soil, animal feed, water, air, used bedding, grass, dung and equipment only 1.2 % were capable of producing cereulide, whereas of the 144 isolates originating from foods 24 % were cereulide producers. Cereulide was detected by two methods: by its toxicity towards mammalian cells (sperm assay) and by its peculiar chemical structure using liquid-chromatograph-mass spectrometry equipment. B. cereus is known as one of the most frequent bacteria occurring in food. Most foods contain more than one kind of B. cereus. When randomly selected 100 isolates of B. cereus from commercial infant foods (dry formulas) were tested, 11% of these produced cereulide. Considering a frequent content of 103 to 104 cfu (colony forming units) of B. cereus per gram of infant food formula (dry), it appears likely that most servings (200 ml, 30 g of the powder reconstituted with water) may contain cereulide producers. When a reconstituted infant formula was inoculated with >105 cfu of cereulide producing B. cereus per ml and left at room temperature, cereulide accumulated to food poisoning levels (> 0.1 mg of cereulide per serving) within 24 hours. Paradoxically, the amount of cereulide (per g of food) increased 10 to 50 fold when the food was diluted 4 - 15 fold with water. The amount of the produced cereulide strongly depended on the composition of the formula: most toxin was formed in formulas with cereals mixed with milk, and least toxin in formulas based on milk only. In spite of the aggressive cleaning practices executed by the modern dairy industry, certain genotypes of B. cereus appear to colonise the silos tanks. In this thesis four strategies to explain their survival of their spores in dairy silos were identified. First, high survival (log 15 min kill ≤ 1.5) in the hot alkaline (pH >13) wash liquid, used at the dairies for cleaning-in-place. Second, efficient adherence of the spores to stainless steel from cold water. Third, a cereulide producing group with spores characterized by slow germination in rich medium and well preserved viability when exposed to heating at 90 ºC. Fourth, spores capable of germinating at 8 ºC and possessing the psychrotolerance gene, cspA. There were indications that spores highly resistant to hot 1% sodium hydroxide may be effectively inactivated by hot 0.9% nitric acid. Eight out of the 14 dairy silo tank isolates possessing hot alkali resistant spores were capable of germinating and forming biofilm in whole milk, not previously reported for B. cereus. In this thesis it was shown that cereulide producing B. cereus was capable of inhibiting the growth of cereulide non-producing B. cereus occurring in the same food. This phenomenon, called antagonism, has long been known to exist between B. cereus and other microbial species, e.g. various species of Bacillus, gram-negative bacteria and plant pathogenic fungi. In this thesis intra-species antagonism of B. cereus was shown for the first time. This brother-killing did not depend on the cereulide molecule, also some of the cereulide non-producers were potent antagonists. Interestingly, the antagonistic clades were most frequently found in isolates from food implicated with human illness. The antagonistic property was therefore proposed in this thesis as a novel virulence factor that increases the human morbidity of the species B. cereus, in particular of the cereulide producers.

Occurrence and properties of steryl ferulates and glycosides in wheat and rye

Cereal kernels are known to contain a number of minor components that possess beneficial health attributes. In this thesis rye and wheat were studied as sources of steryl ferulates and steryl glycosides and their behaviour in processing were evaluated. Further, enzymatic hydrolysis of these conjugates was studied, as well as the capacity of steryl ferulates to inhibit lipid oxidation at different temperatures. Steryl ferulates were shown to have a strong positive correlation with dietary fibre contents in milling fractions from the outer parts of the kernels obtained from a commercial scale mill. Highest contents of steryl ferulates were found in the bran in both cereals, with the content decreasing once moving towards the inner parts of the kernel. Variation in the contents of steryl ferulates was higher in wheat fractions than rye fractions. Steryl glycosides, on the other hand, had either negative or no correlation with dietary fibre, and the range of the steryl glycoside contents was much narrower than that of steryl ferulates in both cereals. There were significant differences in the sterol compositions of these steryl conjugates when compared with each other or with the total plant sterols in the corresponding fractions. Properties of steryl ferulates and steryl glycosides were evaluated after common processing methods and in enzymatic hydrolysis. Thermal and mechanical processing had only minor or no effects on the contents of steryl conjugates from rye and wheat bran. Enzymatic treatments on the other hand caused some changes, especially in the contents of glycosylated sterols. When steryl ferulates extracted from rye or wheat bran were subjected to enzymatic treatments by steryl esterase, significant differences in the rates of hydrolysis were observed between steryl ferulates from different sources with differing sterol compositions. Further, differences were also observed between enzymes from different sources. Steryl glycosides were shown to be hydrolysed by β-glucosidase (cellobiase) from A. niger, but less with β-glucosidases from other sources. Steryl ferulates showed good antioxidant activity at both moderate and high temperatures. In bulk and emulsion systems of methyl linoleate at 40°C steryl ferulates extracted from rye and wheat bran inhibited hydroperoxide formation much more effectively than synthetic steryl ferulates or those extracted from rice (γ-oryzanol), demonstrating that the sterol composition has an effect on the activity. At cooking (100°C) and frying temperatures (180°C) sitostanyl ferulate was shown to inhibit polymer formation significantly and, especially at 100°C, comparably to α-tocopherol. The rate of antioxidant degradation was slower for sitostanyl ferulate, showing higher heat stability than α-tocopherol. When evaluated as a mixture, no synergistic effect was observed between these two antioxidants. The data presented in this thesis provides information that may henceforth be applied when evaluating the intakes of steryl conjugates from cereal sources, as well as their possible influences as minor bioactive components. Wheat and rye both are good sources of steryl ferulates and steryl glycosides and, especially with steryl ferulates, what may be lost out to some other cereals on quantity is compensated with quality of the sterol composition.

Ammonia emissions from pig and cattle slurry in the field and utilization of slurry nitrogen in crop production

Volatilization of ammonia (NH3) from animal manure is a major pathway for nitrogen (N) losses that cause eutrophication, acidification, and other environmental hazards. In this study, the effect of alternative techniques of manure treatment (aeration, separation, addition of peat) and application (broadcast spreading, band spreading, injection, incorporation by harrowing) on ammonia emissions in the field and on nitrogen uptake by ley or cereals was studied. The effect of a mixture of slurry and peat on soil properties was also investigated. The aim of this study was to find ways to improve the utilization of manure nitrogen and reduce its release to the environment. Injection into the soil or incorporation by harrowing clearly reduced ammonia volatilization from slurry more than did the surface application onto a smaller area by band spreading or reduction of the dry matter of slurry by aeration or separation. Surface application showed low ammonia volatilization, when pig slurry was applied to tilled bare clay soil or to spring wheat stands in early growth stages. Apparently, the properties of both slurry and soil enabled the rapid infiltration and absorption of slurry and its ammoniacal nitrogen by the soil. On ley, however, surface-applied cattle slurry lost about half of its ammoniacal nitrogen. The volatilization of ammonia from surface-applied peat manure was slow, but proceeded over a long period of time. After rain or irrigation, the peat manure layer on the soil surface retarded evaporation. Incorporation was less important for the fertilizer effect of peat manure than for pig slurry, but both manures were more effective when incorporated. Peat manure applications increase soil organic matter content and aggregate stability. Stubble mulch tillage hastens the effect in surface soil compared with ploughing. The apparent recovery of ammoniacal manure nitrogen in crop yield was higher with injection and incorporation than with surface applications. This was the case for leys as well as for spring cereals, even though ammonia losses from manures applied to cereals were relatively low with surface applications as well. The ammoniacal nitrogen of surface-applied slurry was obviously adsorbed by the very surface soil and remained mostly unavailable to plant roots in the dry soil. Supplementing manures with inorganic fertilizer nitrogen, which adds plant-available nitrogen to the soil at the start of growth, increased the overall recovery of applied nitrogen in crop yields.

Nitrogen dynamics of organic farming in a crop rotation based on red clover (Trifolium pratense) leys

In agricultural systems which rely on organic sources of nitrogen (N), of which the primary source is biological N fixation (BNF), it is extremely important to use N as efficiently as possible with minimal losses to the environment. The amount of N through BNF should be maximised and the availability of the residual N after legumes should be synchronised to the subsequent plant needs in the crop rotation. Six field experiments in three locations in Finland were conducted in 1994-2006 to determine the productivity and amount of BNF in red clover-grass leys of different ages. The residual effects of the leys for subsequent cereals as well as the N leaching risk were studied by field measurements and by simulation using the CoupModel. N use efficiency (NUE) and N balances were also calculated. The yields of red clover-grass leys were highest in the two-year-old leys (6 700 kg ha-1) under study, but the differences between 2- and 3-year old leys were not high in most cases. BNF (90 kg ha-1 in harvested biomass) correlated strongly with red clover dry matter yield, as the proportion of red clover N derived from the atmosphere (> 85%) was high in our conditions of organically farmed field with low soil mineral N. A red clover content of over 40% in dry matter is targeted to avoid negative N-balances and to gain N for the subsequent crop. Surprisingly, the leys had no significant effect on the yields and N uptake of the two subsequent cereals (winter rye or spring wheat, followed by spring oats). On the other hand, yield and C:N of leys, as well as BNF-N and total-N incorporated into the soil influenced on subsequent cereal yields. NUE of cereals from incorporated ley crop residues was rather high, varying from 30% to 80% (mean 48%). The mineral N content of soil in the profile of 0-90 cm was low, mainly 15-30 kg ha-1. Simulation of N dynamics by CoupModel functioned satisfactorily and is considered a useful tool to estimate N flows in cropping systems relying on organic N sources. Understanding the long-term influence of cultivation history and soil properties on N dynamics remains to be a challenge to further research.

Intensity, productivity and efficiency in agriculture in Finland and implications for N and P fertiliser management

The purpose of this study was to evaluate intensity, productivity and efficiency in agriculture in Finland and show implications for N and P fertiliser management. Environmental concerns relating to agricultural production have been and still are focused on arguments about policies that affect agriculture. These policies constrain production while demand for agricultural products such as food, fibre and energy continuously increase. Therefore the importance of increasing productivity is a great challenge to agriculture. Over the last decades producers have experienced several large changes in the production environment such as the policy reform when Finland joined the EU 1995. Other and market changes occurred with the further EU enlargement with neighbouring countries in 2005 and with the decoupling of supports over the 2006-2007 period. Decreasing prices a decreased number of farmers and decreased profitability in agricultural production have resulted from these changes and constraints and of technological development. It is known that the accession to the EU 1995 would herald changes in agriculture. Especially of interest was how the sudden changes in prices of commodities on especially those of cereals, decreased by 60%, would influence agricultural production. The knowledge of properties of the production function increased in importance as a consequence of price changes. A research on the economic instruments to regulate productions was carried out and combined with earlier studies in paper V. In paper I the objective was to compare two different technologies, the conventional farming and the organic farming, determine differences in productivity and technical efficiency. In addition input specific or environmental efficiencies were analysed. The heterogeneity of agricultural soils and its implications were analysed in article II. In study III the determinants of technical inefficiency were analysed. The aspects and possible effects of the instability in policies due to a partial decoupling of production factors and products were studied in paper IV. Consequently connection between technical efficiency based on the turnover and the sales return was analysed in this study. Simple economic instruments such as fertiliser taxes have a direct effect on fertiliser consumption and indirectly increase the value of organic fertilisers. However, fertiliser taxes, do not fully address the N and P management problems adequately and are therefore not suitable for nutrient management improvements in general. Productivity of organic farms is lower on average than conventional farms and the difference increases when looking at selling returns only. The organic sector needs more research and development on productivity. Livestock density in organic farming increases productivity, however, there is an upper limit to livestock densities on organic farms and therefore nutrient on organic farms are also limited. Soil factors affects phosphorous and nitrogen efficiency. Soils like sand and silt have lower input specific overall efficiency for nutrients N and P. Special attention is needed for the management on these soils. Clay soils and soils with moderate clay content have higher efficiency. Soil heterogeneity is cause for an unavoidable inefficiency in agriculture.

Spatial and temporal determinants of Finnish farmland bird populations

During the past decades agricultural intensification has caused dramatic population declines in a wide range of taxa related to farmland habitats, including farmland birds. In this thesis, I studied how boreal farmland landscape characteristics and agricultural land use affect the abundance and diversity of farmland birds using extensive field data collected by territory mapping of breeding farmland birds in various parts of Finland. My results show that the area and openness of agricultural areas are key determinants of farmland bird abundance and distribution. A landscape composition with enough open farmland combined with key habitats such as farmyards and wetland is likely to provide essential prerequisites for the occurrence of a rich farmland avifauna. In Finland, the majority of large areas suitable for open habitat specialists are located in southern and western parts of the country. However, the diversity of the species with an unfavourable conservation status in Europe (SPECs) had notable hotspot areas in northern and north-western agricultural areas. I found that in boreal agroecosystems farmland birds favour fields with springtime vegetative cover, especially agricultural grasslands and set-asides. Hence, in the spring cereal dominated Finnish agroecosystems it is the absence of field vegetation that may limit populations of many farmland bird species. It is likely that the decrease of crops providing vegetative cover in the spring, such as permanent grasslands, cultivated grass, and autumn-sown cereals, has greatly contributed to the declines of Finnish farmland birds. Grass crops have persistently declined in Finland as a consequence of specialization in crop production and the large-scale decline in livestock husbandry. Small-scale non-crop habitats, especially ditches and ditch margins, are also important for many bird species in the Finnish agroecosystems, but have dramatically declined during the last decades. A major problem for farmland bird conservation in Finland is the conflict between landscape structure and agricultural management. Areas with mixed and cattle farming are virtually absent from the large agricultural plains of southern and south-western Finland, where the landscape structure is more likely to be favourable for rich farmland bird assemblages. On the other hand, mixed and cattle farming is still rather frequent in northern and central parts of the country, where the landscape structure is not suitable for many farmland specialist birds requiring open landscapes. My results provide useful guidelines for farmland bird conservation, and imply that considerable attention needs to be paid to landscape factors when selecting areas for various conservational management actions, such as agri-environment schemes. Actions promoting the abundance of set-asides, grass crops, and ditches would markedly benefit Finnish farmland bird populations. Organic farming may benefit farmland birds, but it is not clear how general its beneficial effect is in boreal agroecosystems. The most urgent action aiming to preserve farmland biodiversity would be to support re-introducing and sustaining cattle farming by environmental subsidies. This would be especially beneficial in the southern parts of Finland, where the landscape characteristics and abundance of agricultural areas are most suitable for farmland birds and where cattle farming is currently rare.

Cereal alkylresorcinols as dietary biomarkers-absorption and occurrence in biological membranes

Several studies link the consumption of whole-grain products to a lowered risk of chronic diseases, such as certain types of cancer, type II diabetes, and cardiovascular diseases. However, the final conclusions of the exact protective mechanisms remain unclear, partly due to a lack of a suitable biomarker for the whole-grain cereals intake. Alkylresorcinols (AR) are phenolic lipids abundant in the outer parts of wheat and rye grains usually with homologues of C15:0- C25:0 alkyl chains, and are suggested to function as whole-grain biomarkers. Mammalian lignan enterolactone has also previously been studied as a potential whole-grain biomarker. In the present work a quantified gas chromatography-mass spectrometry method for the analysis of AR in plasma, erythrocytes, and lipoproteins was developed. The method was used to determine human and pig plasma AR concentrations after the intake of whole-grain wheat and rye products compared to low-fibre wheat bread diets to assess the usability of AR as biomarkers of whole-grain intake. AR plasma concentrations were compared to serum ENL concentrations. AR absorption and elimination kinetics were investigated in a pig model. AR occurrence in human erythrocyte membranes and plasma lipoproteins were determined, and the distribution of AR in blood was evaluated. Plasma AR seem to be absorbed via the lymphatic system from the small intestine, like many other lipophilic compounds. Their apparent elimination half-life is relatively short and is similar to that of tocopherols, which have a similar chemical structure. Plasma AR concentrations increased significantly after a one- to eight-week intake of whole-grain wheat and further on with whole-grain rye bread. The concentrations were also higher after habitual Finnish diet compared to diet with low-fibre bread. Inter-individual variation after a one-week intake of the same amount of bread was high, but the mean plasma AR concentrations increased with increasing AR intake. AR are incorporated into erythrocyte membranes and plasma lipoproteins, and VLDL and HDL were the main AR carriers in human plasma. Based on these studies, plasma AR could function as specific biomarkers of dietary whole-grain products. AR are exclusively found in whole-grains and are more suitable as specific biomarkers of whole-grain intake than previously investigated mammalian lignan enterolactone, that is formed from several plants other than cereals in the diet. Plasma AR C17:0/C21:0 -ratio could distinguish whether whole-grain products in the diet are mainly wheat or rye. AR could be used in epidemiological studies to determine whole-grain intake and to better assess the role of whole-grains in disease prevention.

Charred seeds in Mikkeli Orijärvi : A study of subsistence strategies of an Iron Age settlement in East Finland

In this study the over 350 macrofossil samples, containing over 2300 charred plant remains from an Iron Age settlement containing fossil fields in Mikkeli Orijärvi Kihlinpelto, were studied archaeobotanically. The aim was to get more information about subsistence strategies, especially agriculture and study differences in the plant combinations in the different structures and use the archaeobotanical theory to interpret these structures. The methodological question was to study the taphonomy of the charred plant material. The results gave a diverse impression of the agriculture and subsistence strategies of the settlement in Orijärvi, where barley was the most important cereal with rye, wheat and oat cultivated as minor crops. The arable weed assemblage indicates that the fields were situated in different kinds of soils and the crops were cultivated when different kind of weather conditions were prevailing. Ergot was found with the cereals, and it was growing on some of the arable crops and it also indicates wet climate. Hemp and flax were cultivated and wild plants were collected. The meadow and wetland plants found in the material derive most probably from animal fodder. Tubers of bulbous oat-grass were interesting, because they are usually found in graves. Comparison with other Iron Age settlements and graves indicates that the plant material found from the ancient field layers derives most probably from dwellings and graves, which were taken into cultivation.