Crop & Food Research Confidential Report No. 143 Nutrition and health qualities of potatoes - a future focus C E Lister & J Munro March 2000 A report prepared for New Zealand Federation of Vegetable and Potato Growers Copy 13 of 13 New Zealand Institute for Crop & Food Research Limited Private Bag 4704, Christchurch, New Zealand Circulation of this report is restricted. Consult the author(s) and the institute’s scientific editor about obtaining further copies. This report may not be copied in part or full. © 2000 New Zealand Institute for Crop & Food Research Limited Contents 1 Executive summary 1 2 Potatoes and their consumption 2 2.1 3 1997 National Nutrition Survey The composition of potatoes 3 4 3.1 Dry matter and water content 5 3.2 Carbohydrates 5 3.2.1 3.2.2 3.2.3 5 6 7 3.3 Starch Sugars Dietary fibre (non-starch polysaccharides) Nitrogen compounds 3.3.1 3.3.2 3.3.3 3.3.4 Protein Free amino acids Inorganic nitrogen Glycoalkaloids 8 8 10 10 11 3.4 Lipids 11 3.5 Vitamins 11 3.6 Minerals 13 3.7 Phenolic compounds 14 3.7.1 3.7.2 3.7.3 3.7.4 3.7.5 3.7.6 14 15 15 16 16 16 3.8 3.9 Role of phenolics in browning reactions Antioxidant activity Antimutagenic and anticarcinogenic effects Glucose-lowering properties Cholesterol-lowering effect Effect on protein nutritional quality Plant pigments 17 3.8.1 3.8.2 3.8.3 17 17 18 Carotenoids Anthocyanins Others Nutrient losses/changes during cooking 18 4 The health benefits of potatoes 19 5 The future 21 5.1 5.2 Feeding the world in the twenty-first century 21 5.1.1 5.1.2 5.1.3 5.1.4 Protein quality improvement Provitamin A Disease and pest resistance Minerals 22 22 23 23 Delivery of additional health benefits 24 5.2.1 5.2.2 24 26 Increased levels of phenolics and other antioxidants Recombinant proteins/vaccines DCom4:Users:dcom4:Desktop:NZ HORT REPORTS:C & F potatoes143.doc 5.2.3 5.2.4 5.3 Modified carbohydrate Other 27 27 Removal of antinutritional compounds and undesirable chemicals 27 5.3.1 5.3.2 27 28 Glycoalkaloids Chemical residues 5.4 Browning prevention 28 5.5 Other new and improved potato products 29 5.5.1 5.5.2 29 29 5.6 “Healthier” fried products Novelty coloured potato products Utilizing potato waste 30 6 Recommendations/future research 30 7 Acknowledgements 31 8 References 32 Appendix I - the chemical composition of New Zealandgrown potatoes, raw and cooked by a variety of method DCom4:Users:dcom4:Desktop:NZ HORT REPORTS:C & F potatoes143.doc 39 1 Executive summary This report describes the results of a literature review undertaken for the potato sector of Vegfed to describe the nutritional status and health benefits of potatoes. It highlights current knowledge and identifies some prospects for the future. It also explores the next generation of key attributes that may be used for product development and in marketing strategies. The key points are noted below. ♣ Potatoes are a vegetable whose nutritional value is underestimated. They are often believed to be a high energy food that provides little in the way of nutrients. However, potatoes are a considerably richer source of nutrients than of energy. In brief, potatoes are: Potatoes are a vegetable whose - a source of several vitamins, especially vitamin C and some important B group vitamins, - rich in minerals such as potassium and iron, - a source of phenolics, compounds that may have an important role in health, - virtually free of fat, although they are easily turned into fatty foods, - almost free of soluble sugar, that provides little in - of low energy density - they ‘fill you up’ without providing many calories, the way of nutrients. - a source of high quality protein, although they are deficient in the essential amino acid methionine, - readily digested but they also have a high water content so weight for weight there is a relatively low impact on blood sugar. ♣ On the other side of the ledger, potatoes may also contain glycoalkaloids (often found in green tubers) although the risk that they contain toxic levels is minimal. ♣ Despite potatoes already having a number of desirable nutritional qualities there is still considerable scope to improve the composition of potatoes in a number of areas. These improvements may be achieved through traditional plant breeding, genetic modification, and/or changes in agronomic and postharvest handling procedures. They include: - boosting nutritional quality by increasing the levels of essential amino acids, β-carotene (a precursor of vitamin A) or other antioxidants, - removing or limiting the levels of glycoalkaloids, Nutrition and health qualities of potatoes – a future focus C Lister, March 2000 Crop & Food Research Confidential Report No. 143 New Zealand Institute for Crop & Food Research Ltd Page 1 nutritional value is underestimated. They are often believed to be a high energy food This is simply not true. 2 - developing new cultivars that absorb less fat during frying, - using potatoes to produce genetically engineered vaccines and other high value pharmaceutical products, - developing novelty coloured potato products that have added health benefits. ♣ Initially new potato cultivars could be developed followed by new potato products tailored to meet specific nutritional requirements of certain sectors of the population. ♣ As new cultivars are developed with additional health benefits there will be a need to re-evaluate the potential of potato wastes to better utilize key components with the aim of improving human or animal health. Potatoes and their consumption The potato is an extremely important food crop. It is grown in more countries than any other crop except maize. Its volume of production ranks fourth in the world after rice, wheat and maize. Potatoes are number one in terms of household expenditure on vegetables in New Zealand. In 1998 the total household expenditure on potatoes was $65.07 million, up from $45.8 million in 1984 (Commercial Grower, March 1999). Daily consumption of potatoes per capita varies considerably and depends on age, sex, eating habits and daily activities of consumers. In most countries potato consumption is not influenced by family income. US per capita consumption of potatoes is about 61 kg/year, for the UK it is 105 kg/year, for the USSR 110 kg/year and it may even reach close to 150 kg/year in Poland and Germany (Lisiska & Leszcyski 1989). Figures from the recent National Nutrition Survey (Russell et al. 1999) show that New Zealand consumers of potatoes eat an average of 180 g of potatoes (in all forms) per day. For females the average is 140 g per day while for males it is 219 g per day. Thus, potatoes are a very significant part of the diet in many countries and can make a significant contribution to human nutrition. The global production of potatoes has been estimated at 350 million tones per annum (Friedman & McDonald 1997). New Zealand’s share is approximately 490 000 tonnes per annum, some of which is used for domestic seed (Table 1). The countries with the largest production volumes of potatoes are the USSR, China, Poland, US and Germany (Lisiska & Leszcyski 1989). In addition to human consumption, a large proportion of potatoes are used as stock feed. The nutritional value of potatoes is also important. The higher the nutritional quality the better the stock feed, lessening the requirement for potatoes to be mixed with other higher value feeds. Considerable amounts of potatoes are used for processing. Volumes have been increasing rapidly in the last couple Page 2 of decades. Nutritional considerations are important when processing potatoes. Some of the issues are different from those for the fresh market. Table 1: Potato production in New Zealand (combined domestic and export use), volume in tonnes (from Heyes et al. 1997) Fresh Seed Process Stockfeed 2.1 1995/6 259 000 30 000 200 000 1983/4 194 305 29 262 60 938 10 086 1997 National Nutrition Survey In 1997 a New Zealand National Nutrition Survey (NNS97) was funded by the Ministry of Health and conducted by the LINZ® Activity & Health Research Unit of the University of Otago. The aim of this survey was to collect information on food and nutrient intakes, dietary habits and nutrition-related clinical measures of New Zealanders. NNS97 was based on a nationally representative sample of 4636 New Zealanders living in selected households and aged 15 years or more. Some of the data from the survey are summarised below (Russell et al. 1999). Note that in most cases potatoes also included kumara. ϒ Potatoes are the most commonly consumed vegetable in the New Zealand diet; 95% of males and 93% of females ate potatoes (boiled, mashed, baked or roasted) at least once a week (for kumara the pattern was 27% and 32% for males and females respectively) while 54% of males and 42% of females ate hot potato or kumara chips, French fries or wedges at least once a week. ϒ Older people are more frequent consumers of potato. Provincial people ate potatoes more often than people living in metropolitan areas. Rice and pasta consumption showed the reverse trends. ϒ 7% of energy in the diet was contributed by potatoes and kumara. ϒ 6% of total fat was contributed from potatoes and kumara, for young males (19-24 years) the contribution was 10%. ϒ Potatoes and kumara were a principal carbohydrate source, contributing 10% of total carbohydrates, this was higher for males (11%) than females (9%). ϒ Vegetables (including potato and kumara) were the greatest source of dietary fibre in the diet at 28%. For potatoes and kumara alone the figure was 11%. Again the figures were higher for males (13%) than females (10%). ϒ Potatoes and kumara provided 7% of daily iron intake, 8% of folate, 4% of protein and 2% of calcium. Page 3 In terms of the general health of the New Zealand population, 17% of New Zealanders are regarded as obese (that is a body mass index of greater than 32 for New Zealand M∼ori and Pacific people and greater than 30 for New Zealand European and others) and an additional 35% were considered overweight. The obesity rate has risen from 11% in 1989 for both males and females. About 22% of males and 18% of females had high blood pressure. However, the mean serum cholesterol in NNS97 (5.7 mmol/L) has decreased slightly from the 1989 level (5.9 mmol/L), although 23% of the population still has serum cholesterol greater than 6.5 mmol/L. These figures indicate that there is considerable room for improvement in the health of New Zealanders. Diet is one major way to improve health. 3 The composition of potatoes Numerous books and papers cover the composition of potatoes (for example Burton 1989; Lisiska & Leszcyski 1989). Table 2 shows the range of chemical composition of potato tubers using data from around the world. Data on the composition of New Zealand-grown potatoes are available in the New Zealand Food Composition Database, and some of these data are presented in Tables 8-12 in Appendix I. For comparison, the composition of other foods that may replace potatoes in a meal (e.g. rice, pasta and bread) are also provided. There is a considerable range in the concentration of each Table 2: The typical chemical composition of potato tubers (taken from Lisiska & Leszcyski 1989). Content (%) Substance Dry matter Starch Reducing sugars Total sugar Crude fibre Pectic substances Total nitrogen Crude protein (total nitrogen x 6.25) Protein nitrogen in total nitrogen Amide nitrogen Amino acid nitrogen Nitrates Lipids Ash Organic acids Ascorbic acid and dehydroascorbic acid1 Glycoalkaloids1 Phenolic compounds1 1 In mg/100 g. Page 4 Range Mean 13.1–36.8 8.0-29.4 0.0-5.0 0.05-8.0 0.17-3.48 0.2-1.5 0.11-0.74 0.69-4.63 27.3-73.4 0.029-0.052 0.065-0.098 0.0-0.05 0.02-0.2 0.44-1.87 0.4-1.0 1-54 0.2-41 5-30 23.7 17.5 0.3 0.5 0.71 0.32 2.00 54.7 0.12 1.1 0.6 10-25 3-10 - of the components. The chemical composition of potato tubers is mainly controlled by genetic factors. In addition, the composition is affected by the age and maturity of the tubers as well as environmental conditions, i.e. climate, soil and cultural practices. The composition of potato tubers also changes during storage and is affected by the way in which they are processed. The significant potato components, in terms of health benefits, are discussed in more detail below. 3.1 Dry matter and water content Dry matter content of potato tubers ranges from about 13 to 37% with an average of 24% (Table 2). The other 75% of the potato consists of water. Dry matter content increases during the growing season and is highest in the vascular system, intermediate in the cortex and lowest in the pith. The dry mater or ‘solids content’ of tubers is one of the prime characters used by potato processors to evaluate a crop. Potatoes with high dry matter are most suitable for the manufacture of dehydrated food products and stockfeed and are especially good for the production of fried foods. The dry matter of potato tubers is composed of a number of substances that are either soluble or insoluble in water. Dry matter is particularly significant when frying as the greater the dry matter the less fat uptake because there is less water for it to replace. Table 3 shows the average contents of potato tuber dry matter. There may be considerable changes in dry matter after cooking potato, particularly frying (see Table 8 in Appendix I). Table 3: Components or dry matter of potato tuber (taken from Lisiska & Leszcyski 1989). Component Starch Total sugars Crude fibre Crude protein Crude lipids Ash 3.2 Mean content (%) 75.30 2.10 2.32 7.94 0.50 4.41 Carbohydrates Carbohydrates make up the greatest proportion of the dry weight of potato tubers (Table 3). Carbohydrates are present in the form of starch, sugars and non-starch polysaccharides (cell wall components, often termed dietary fibre). Available carbohydrate values for New Zealand-grown potatoes are given in Table 8 in Appendix I. 3.2.1 Starch Starch is a major component of the potato tuber at approximately 17 g/100 g FW or 75% of its dry weight (Tables 2 and 3). During the growing season starch accumulates in the cells of tubers, forming single or complex granules. Starch, being a carbohydrate, is an important energy source and can affect quality. Potato starch is an important part of food products and is a raw Page 5 material for industry. Starch occurs in a number of forms that differ in molecular structure and their susceptibility to digestion. Starch may be classified into digestible starch (DS) and resistant starch (RS). From a health standpoint, susceptibility to digestion is important for several reasons. 1. Starch is a chain of glucose molecules, and during digestion starch chains are typically broken down very quickly in the small intestine into maltose and then glucose. Starch digestion can, therefore, have a large impact on blood glucose levels, which need to be closely controlled in people with diabetes. 2. Starch that cannot be digested before passing from the small intestine is fermented by bacteria in the colon, with the production of short chain fatty acids. The short chain fatty acids from starch fermentation provide the body with about half as much energy as would have been obtained from the starch if it had been digested in the small intestine. 3. There is increasing scientific evidence that resistant starch can act as a prebiotic, increasing the population of beneficial bacteria, such as bifidobacteria, and having a beneficial effect on the ecology of the large bowel (Bird 1999). Before consumption, starch must be gelatinised. Consumption of ungelatinised potato starch reduces protein digestibility and induces toxicity (Lisiska & Leszcyski 1989). Resistant starch (RS) in raw potatoes is high. However, different RS values were obtained when tubers were processed, ranging from 1.18% in boiled potatoes to 10.38% in retrograded flour (GarciaAlonso & Goni 2000). The starch in cooked potatoes is generally easily digested, but more so in mature than in new potatoes. In a recent study comparing three potato varieties, four cooking methods, and two states of maturity, the impact of potatoes on blood glucose levels, reflected in the glycaemic index, was exceptionally high, regardless of variety, cooking method and maturity (Soh & Brand-Miller 1999). New potatoes had lower GI values than mature potatoes, probably because, with maturation, the amylose component of starch becomes increasingly branched, and thus increasingly susceptible to digestion. In cold cooked potato and mashed potato, starch digestibility is reduced due to its retrogradation. Repeated cooling and mashing increases the amount of starch that is indigestible by humans (Englyst & Cummings 1987). 3.2.2 Sugars The sugar content of potatoes is very low at an average of 0.5% of the wet weight or just over 2% of dry weight (Tables 2 and 3). However, sugar content is highly variable, depending on the type, maturity and physiological state of the potato. In free form the following sugars are found in potato tubers: the monosaccharides glucose and fructose, which are reducing sugars, and sucrose, a non-reducing disaccharide. A high sugar content (especially reducing sugars) renders potato tubers unsuitable for use as raw material for processing, especially for dehydrated and fried products. Page 6 3.2.3 Dietary fibre (non-starch polysaccharides) Dietary fibre consists mainly of the polysaccharides of plant cell walls. The dietary fibre may “dilute” highly caloric components in food, stimulate peristaltic movement and retard digestion of some other food components. The essential non-starch polysaccharides in potatoes are cellulose, hemicellulose, pentosans and pectic substances. Starch is also a polysaccharide, but the sugars and the way they are bonded to one another in cell walls and in starch are different. The human gut produces enzymes to break down the structure of starch, but they cannot attack cell wall polysaccharides, which therefore survive the small intestine as non-digested “dietary fibre”. Potatoes contain a relatively small proportion of dietary fibre because they are a storage organ in which starch becomes the dominant constituent in the mature tuber. Most potatoes contain about 1-3% dietary fibre compared with about 17% starch, on an edible weight basis (Table 2 and New Zealand data in Table 8 in Appendix I). Potatoes consist of a mass of thin-walled cells full of starch. As well as being thin, the cell walls of potatoes are not lignified. Lignin is a resistant material that encrusts cell walls such as those found in wheat bran, making them resistant to fermentation in the large bowel and an excellent faecal bulking agent. Being thin, and not lignified, the cell walls of potatoes are almost totally fermented by bacteria in the colon. The dietary fibre in potatoes thus has almost no effect on faecal bulk and does little to prevent constipation. However, during fermentation of dietary fibre in the colon short chain fatty acids are produced. One of the short chain fatty acids, butyric acid, is thought to provide some protection against colorectal cancer by inducing apoptosis, which is the destruction of abnormal cells. Butyric acid would be most useful if it were produced where most of the colorectal cancer occurs, that is, in the distal colon. However, potato cell walls are so thin and fermentable that most of the butyric acid produced is likely to be in the proximal rather than the distal colon. The cell walls could be induced to ferment in the distal colon only if there was some other bulky non-digested material in the diet that increased movement of gut contents sufficiently for the potato fibre to reach the distal colon before being fermented. One portion of the cell wall fraction of potatoes that might have a more positive impact on health is the potato peel fraction, not because of the cell walls per se but because they are impregnated with materials that provide the potato with a resistant coating. The resistant, water proofing material of the potato “coat” consists largely of suberin, which, because it is non-polar, has the capacity to bind some of the most potent food carcinogens, the heterocyclic amines (Harris 1999). Heterocyclic amines are produced during cooking and charring of proteins, as occurs abundantly during barbecuing. So eating a whole baked potato with barbequed food may help “mop-up” some of those carcinogens. Other components of potato peel, such as phenolics, may also have beneficial effects on human health (see Section 3.7). Page 7 3.3 Nitrogen compounds Nitrogen compounds are the second major component in potato tubers, after carbohydrates. Their amount in conversion to total protein (N x 6.25) ranges from 2.77 to 14.6% of dry weight (Table 3 and New Zealand data in Table 8 in Appendix I). The content of total nitrogen increases upon maturation. Approximately 90% of the nitrogen present in potato tubers is in the form of compounds that are soluble in water. Insoluble nitrogen compounds are found mostly in the skin. About 50% of the total nitrogen from potatoes is derived from proteins; the remaining nitrogen consists of free amino acids, amide nitrogen associated with asparagines and glutamine, nitrogen of other organic compounds, inorganic nitrogen and alkaloid nitrogen (Lisiska & Leszcyski 1989). 3.3.1 Protein The essential nitrogen fraction in a potato tuber is protein nitrogen. Potatoes are commonly perceived as a carbohydrate source only, but they do contain high quality protein. On a fresh weight basis they contain only about 2% protein; the value increases to about 8% when examined on a dry weight basis (Tables 2 and 3 and New Zealand data in Table 8 in Appendix I). That makes potatoes comparable to cereals, such as rice or wheat (McCay et al. 1987). In countries where potato consumption is high this vegetable can make a significant contribution to health as a protein source (Woolfe 1986). Due to the dietary quality of tuber nitrogen, 100 g of boiled potatoes supplies 8-13% and 6-7% of the FAO-WHO recommended daily allowance of nitrogen for children and adults, respectively (Horton & Sawyer 1985). The contribution of protein nitrogen to total nitrogen depends on potato cultivar, environmental conditions and cultural practices, especially nitrogen fertilization application rates. Proteins are important constituents of cellular membranes as well as various cytoplasmic structures. Also the enzymes present in potatoes are made up of proteins. The fraction of proteins that is not easily soluble is built up in the cell walls. The major fraction of proteins present in potato tubers is constituted of simple proteins. Potato proteins comprise 18-20 amino acids present in varying quantities (Table 4 and Table 11 in Appendix I). Humans cannot synthesize some amino acids and so they must to be provided in the foods we eat. These amino acids are called essential amino acids and include methionine, threonine, tryptophan, valine, lysine, isoleucine, leucine and phenylalanine. Deficiency in any of these amino acids limits the quality of the protein. There are a number of ways to measure the quality of protein contained in different foods. These include the chemical score (CS), which is the ratio of limiting amino acids in the tested protein to amino acid content in standard protein, expressed as a percentage. Protein of the whole hen egg is usually referred to as standard protein. Other indices of protein nutritional value are used on the basis of protein conversion rate such as: (a) protein efficiency ration (PER), which expresses body weight gain ratio to protein intake; (b) net protein utilization (NPU) or (c) biological value (BV) expressed in % of nitrogenous substances absorbed by the digestive tract. Page 8 A summary of the nutritive value of potato protein (Markakis 1975) shows: 1. on the basis of amino acid composition, the calculated protein quality is about 70% that of whole egg protein. This is higher than that of most other food plants, 2. potatoes provide a good source of lysine but low contents of sulphurcontaining amino acids (methionine, cysteine) limit their nutritive value. Tuber methionine is the limiting essential amino acid for human nutrition (Kaldy & Markakis 1972; Rexen 1976), 3. human feeding trials suggest that potato proteins are of a very high quality, possibly higher than indicated by their amino acid composition. This may be because protein utilization is enhanced by the high content of free amino acids and other metabolites mentioned above. Since potatoes need to be baked, boiled, fried, or otherwise cooked before consumption, it is of interest to determine to what extent exposure to heat affects nutritive value of the proteins. Losses during chipping, canning and drum-drying are considerable (Jaswal 1973), but appear to be minor during boiling and frying (Friedman 1992a and b). Heat treatment of potatoes destined for direct consumption (cooking, baking) does not cause significant changes in total nitrogen content or protein content, except from losses on peeling (Toma et al. 1978; Weaver et al. 1983). When potatoes are cooked some loss of free amino acids and combined amino acids occurs. The losses are higher when low specific gravity potato tubers are cooked and they affect non-essential amino acids more than essential amino acids (Jaswal 1973). Protein concentrates prepared from potatoes have excellent nutritional quality when measured in terms of protein efficiency ratio (PER = 2.90), biological value (BV = 79.5), net protein utilization (NPU = 74.2) and nitrogen retention (Nestares et al. 1993). If potato protein is supplemented with protein that is rich in methionine the nutritive value of such a mixture is higher than that of either protein separately. The conversion rate of such a mixture is considerably higher and thus the amount of protein necessary to satisfy daily requirements is lower. The amount of potato protein necessary to satisfy daily requirements is a little higher than that of whole egg but almost the same as the protein present in whole milk and about 40% lower than wheat protein. Potato protein in combination with egg protein (rich in sulphur-containing amino acids, such as methionine) is of high biological value. The combination of these two proteins alone in the proportion 65:35 potato to egg will furnish the daily requirements if supplied to the organism in an amount 30% lower than provided by egg alone (Lisiska & Leszcyski 1989). Thus, potato protein merits inclusion in various food formulations as a source of high quality protein. Table 4: Amino acid composition of potato tubers (adapted from Lisiska & Leszcyski 1989). Amino acid Alanine Composition of amino acids in tuber protein (%) 4.62-5.32 Page 9 Free amino acids (mg/100 g DW) 6-251 Arginine Aspartic acid Cysteine Glutamic acid Glycine Histidine Isoleucine Leucine Lysine Methionine Phenylalanine Proline Serine Threonine Tryptophan Tyrosine Valine 3.3.2 4.74-5.7 11.9-13.91 0.2-1.25 10.23-11.8 4.3-6.05 2.06-2.5 3.73-5.8 9.7-10.28 6.7-10.05 1.2-2.15 4.8-6.53 4.7-7.38 4.9-5.92 4.6-6.5 0.3-1.85 4.5-5.68 4.88-7.4 42-736 123-892 0-9 76-766 1-52 15-328 14-256 8-157 9-319 9-128 0-266 0-484 15-197 14-270 1-174 9-319 15-405 Free amino acids More than two-thirds of the non-protein nitrogen present in potatoes is present as free amino acids. A number of different amino acids have been detected; aspartic acid, glutamic acid and valine are generally present in the highest quantities (Lisiska & Leszcyski 1989). Table 4 shows the free amino acid composition of potato tubers. Tyrosine is one of the free amino acids that has attracted a lot of attention. Tyrosine, oxidized by polyphenol oxidase, changes colour and causes darkening of potato flesh, a process termed enzymic browning (see Section 3.7.1). A close relationship has been reported between tyrosine content and discoloration (Stark et al. 1985), although other researchers have not observed this relationship. 3.3.3 Inorganic nitrogen The inorganic nitrogen found in potato tubers occurs mainly as ammonia compounds, nitrates and sometimes nitrites, although the latter are present only in trace amounts. In recent years some attention has been focused on the nitrate content of potato tubers. As in all food products this is associated with the conversion of nitrates to nitrites in all living organisms. Nitrites subsequently produce carcinogenic nitrosamines. The nitrate content of potato tubers is generally within the range of 4.0 to 25 mg NO3 per 100 g fresh weight. Unfavourable conditions can cause the levels to go even higher. The levels are higher in potato skin than flesh (Lisiska & Leszcyski 1989). The levels are considered to be low and are similar to those in bananas, tomatoes, cucumbers or peas although higher than in some vegetables such as spinach, lettuce and cabbage. Large losses of nitrates occur during washing, peeling and cooking of potatoes. Page 10 3.3.4 Glycoalkaloids Members of the Solanaceae family, and the Solanum genus in particular, synthesize a variety of alkaloidal compounds. The most frequently encountered are the commonly named glycoalkaloids, which are nitrogencontaining steroidal glycosides. Potato glycoalkaloids have been the subject of an extensive recent review (Friedman & McDonald 1997). In commercial potato cultivars the primary compounds are α-solanine and α-chaconine, glycosides of the steroidal alkaloid solanidine. These and similar compounds have toxic effects in humans (Friedman & McDonald 1997). This toxic effect occurs only when glycoalkaloid intake is very high. At sublethal doses (3-5 mg/kg body weight) for adults the amount of glycoalkaloid intake would have to be 200-350 mg. The chances that such a quantity of glycoalkaloid could be consumed in a potato meal are extremely remote. The glycoalkaloids produce a tart astringent taste and larger quantities even impart a sharp bitter taste, so it is a warning against consuming them. The common current guideline for potatoes establishes an upper level of 20 mg/100 g FW of glycoalkaloids. However, the glycoalkaloid content of the majority of potato cultivars is between 3 and 10 mg per 100 g of tubers. The levels of glycoalkaloids present in potatoes may be affected by a variety of factors, including the particular variety, cultural practices and climatic conditions at the growing area (Morris & Petermann 1985). Tubers do not stop producing glycalkaloids after harvest and improper handling and/or storage conditions may cause a dramatic rise in these compounds (Salunkhe et al. 1972). Levels can be especially high in green or damaged potatoes. Processing may also affect glycoalkaloid content. While glycoalkaloids seem to be largely unchanged by cooking, other types of preparation such as slicing and peeling may alter the levels (Friedman & McDonald 1997). This is because the concentrations of glycoalkaloids are highest in the peels and, since they are soluble in water, they are often leached into the cooking liquid. 3.4 Lipids Lipids, also called crude fat, are present at very low levels in raw potatoes (Tables 2 and 3) and they contain no cholesterol. Thus, the energy value provided by potato fats and their effect on obesity are not important. However, frying potatoes, roasting or adding milk and butter during mashing can change this considerably (see details for potatoes cooked by different methods in Table 8 in Appendix I). 3.5 Vitamins Potatoes contain some vitamins in significant amounts and are a much better source than other foods which may replace them in a meal (see Table 10 in Appendix I). Of all the vitamins, potatoes contain vitamin C in the highest quantity (Table 5). Vitamin C is a water-soluble vitamin and its main component is ascorbic acid. However, dehydroascorbic acid has properties similar to vitamin C if it is reduced to ascorbic acid in the organism. The total amount of the two acids in potato tubers ranges from 1 to 54 mg per 100 g Page 11 fresh weight, although most frequently it is between 10 and 25 mg/100 g. The contribution of dehydroascorbic acid to total vitamin C content averages 12-15% (Lisiska & Leszcyski 1989). The highest concentration of vitamin C is found in the vicinity of the vascular system and is lowest in the pith and skin. There has been some breeding of potatoes to increase the levels of vitamin C. There may be considerable losses of vitamin C during processing, and the losses are different for peeled and unpeeled potatoes. Typically, vitamin C losses from potatoes boiled in their skins are 18-24% while peeled boiled potatoes lose 35-50% (Lisiska & Leszcyski 1989). Some of the loss is due to leaching into the cooking water and the rest is decomposed during cooking. Vitamin C losses due to frying reach 75% while baking losses are generally lower at 25-40%. Data from the New Zealand Food Composition tables indicate that microwaving may result in large losses of vitamin C (see Table 10 in Appendix I). However, the amount of vitamin C remaining in potatoes after cooking is still high and it makes a considerable contribution to the recommended daily intake of vitamin C (see Section 4). A typical portion of 150 g of boiled or baked potatoes will provide about 30% of the daily requirement of vitamin C. Potato tubers also contain vitamin B1 (thiamine), but other water-soluble vitamins are present in fairly low amounts (see Table 5 and New Zealand data in Table 10 in Appendix I), although some do make a contribution to the recommended daily intakes of vitamins (see Section 4). There is some loss of these vitamins during processing. Fat-soluble vitamins are present in trace amounts. Provitamin A is contributed by some of the carotenoids. Not all carotenoids have provitamin A activity, the main one is ∃-carotene (for details on carotenoids see Section 3.8.1). Table 5: Vitamin content in potato tubers (taken from Lisiska & Leszcyski 1989). Content (µg/100 g) Vitamin Substance Water soluble C Ascorbic acid + dehydroascorbic acid B1 Thiamine B2 Riboflavin B6 Pyridoxine PP Niacin Pantothenic acid Folic acid H Biotin Fat soluble Provitamin A K1 Phytonadione Page 12 Range Mean 1000-54000 10000-25000 24-180 7-200 9001 360-3300 190-320 5-33 0.6 100 70 1000 - 11-56 60-80 1 - 1 3.6 In dry matter. Minerals Mineral substances, called ashes, average 1.1% in potato tubers (Table 2 and New Zealand data in Table 9 in Appendix I). Potassium is found as the major cation in potato tubers. Higher concentrations of potassium are present in the skin and directly beneath it than in the interior of the potato tuber. Potassium content increases during the entire growing season. Potassium plays an important role in water and ionic supply in humans. In several disease (e.g. renal insufficiency) a potato diet is recommended due to its high potassium content along with other desirable nutritional characteristics. Other minerals present in moderate quantities include phosphorus, chlorine, sulphur and magnesium. Calcium is present in small quantities, mostly in the skin and the vascular system. Iron is also present in fairly low amounts but may make a contribution to dietary intake (see Section 4). Iron, in association with chlorogenic acid, causes after-cooking darkening of potatoes. During preparation of potatoes for consumption a portion of the minerals present in the skins is lost with peeling. Heat treatment on the other hand does not result in significant mineral losses (True et al. 1979). The potato itself is low in sodium, which is desirable. However, the amount of sodium is often increased ten-fold by the addition of salt during or after cooking (Table 9 in Appendix I). Table 6: Composition of potato tuber ash (adapted from Lisiska & Leszcyski 1989). Element Potassium (K) Phosphorus (P) Chlorine (Cl) Sulphur (S) Magnesium (Mg) Sodium (Na) Calcium (Ca) Silicon (Si) Iron (Fe) Aluminium (Al) Manganese (Mn) Zinc (Zn) Copper (Cu) Content in tuber dry matter (mg/100 g DW) 1400-2500 120-600 45-800 40-400 45-220 0-330 10-130 5-89 2.5-72 0.2-35 0.5-8 0.8-2.2 0.06-2.8 Page 13 3.7 Phenolic compounds Potato tubers contain a number of phenolic compounds, but their percentage is rather low (Table 2). Some of them appear in a free form while others are bound. The phenolic compounds present in the tubers include polyphenols, monohydric phenols, coumarins, anthocyanins, flavones, tannins and lignin. Phenolic compounds are distributed mostly between the cortex and skin (peel) tissues of the potato. There are about ten times as much phenolic compounds in the peel as in the flesh of the potato (Lisiska & Leszcyski 1989). Some substances are only detected in the peel. Only tyrosine (a monohydric phenol) is highest in the interior of the tuber and lowest in the external layer. Tannins are mostly localized in the outer layers of the potato, imparting a tan shade to the skin. The chemistry, biochemistry and dietary role of potato polyphenols have been reviewed by (Friedman 1997). About 50% of the polyphenolic compounds are located in the peel and adjoining tissues while the rest decreases in concentration from the outside towards the centre of the potato tuber (Hasegawa et al. 1966). Chlorogenic acid constitutes up to 90% of the total polyphenolic content of potatoes (Friedman 1997). Other phenolic acids include protocatechuic, sinapic, coumaric and vanillic acids. Other polyphenolic compounds present in potatoes include anthocyanins (see Section 3.8.2), flavanones (naringenin and eriodictyol), flavan-3-ols (catechin and epicatechin) and flavonols (kaempferol and sometimes quercetin glycosides) (Lewis 1996). Many of these compounds are present in fairly low concentrations. Tannins are mostly localized in the outer layers of the potato and impart a tan shade to the skin. Polyphenols may have little dietary significance if they are destroyed during processing (baking, boiling, frying, etc.). Destruction of chlorogenic acid is least during microwaving but the fate of phenolics during processing has not been fully studied. For components such as phenolics that are present in much higher concentrations in the peel the amount of peel consumed is important. The peel of baked or fried potatoes is the principal source of peel in the human diet (Bushway et al. 1983; Friedman & Dao 1992). 3.7.1 Role of phenolics in browning reactions Browning reactions are very common in potatoes and potato products, and in some cases are undesirable from a visual point of view (although in other cases they are desirable). Enzyme-catalyzed browning reactions involve the oxidation of phenolic compounds by the enzyme tyrosinase (polyphenol oxidase, PPO) to quinines, followed by transformation of the quinines to dark pigments. Chlorogenic acid seems to be responsible for the bluish-grey discoloration of boiled or steamed potatoes following exposure to air. This so called “after-cooking blackening or darkening,” which can occur within a few minutes after steam peeling, is perceived by many consumers as undesirable. The enzymatic and nonenzymatic browning reactions of amino acids and proteins with carbohydrates, oxidised lipids, and oxidised phenolics will cause food to deteriorate during storage and processing. In addition, a Page 14 loss in nutritional quality and potentially in safety is attributed to the destruction of essential amino acids, a decrease in digestibility, the inhibition of proteolytic and glycolytic enzymes, interaction with metal ions and the formation of antinutritional and toxic compounds. However, some browning reaction products may have beneficial effects, including antimutagenic activity since browning reactions lead to the formation of naturally occurring antioxidants, antibiotic activity and antiallergenic properties (Friedman 1996b). The sometimes green colour of potato cooking water appears to be due to a pigment derived from the reaction of chlorogenic acid and glutamine (Adams 1994). Whether the heat-induced discoloration adversely affects the nutritional value of boiled and darkened potatoes is not known. 3.7.2 Antioxidant activity Polyphenolic compounds in potatoes show antioxidative activity in several systems. In a US study total phenol content of potato (peeled) was 28 mg per 100 g FW and it was ranked twentieth out of 23 commonly consumed vegetables. However, it was ranked ninth in terms of antioxidant activity (Vinson et al. 1998). This study gave potatoes as the fourth greatest contributor to phenolics consumed per day from vegetables. Chlorogenic acid from potato has been found to be an effective inhibitor of lipid oxidation (AlSaikhan et al. 1995). Extracts from potato peels possess strong antioxidant activity, attributed mostly to their chlorogenic, protocatechuic and caffeic acid contents (Onyeneho & Hettiarachchy 1993). Extracts prepared from red peels have stronger activity than those from brown peels, probably due to the strong antioxidant activity of the anthocyanins. New Zealand studies (Lister & Podivinsky 1998; Lister 1999; Lister et al. 2000) have measured the antioxidant activity of New Zealand-grown potatoes. Red Desiree ranked ninth out of 15 vegetables and Rua was eleventh. Antioxidant activity was correlated with the phenolic content of the vegetables, and the two potato cultivars ranked eighth and tenth respectively when vegetables were compared on an equal weight basis. If the weight of potato consumed per day is taken into consideration then potatoes move up the rankings and are better than many vegetables. However, compared to other rich sources of antioxidants (e.g. red wine, berry fruit) potatoes are a small contributor to the potential daily intake of antioxidants. Peeling the potato considerably reduced both the phenolic content and the antioxidant activity. Other potato cultivars have higher phenolic contents (Lewis 1996) and thus may have higher antioxidant activity. 3.7.3 Antimutagenic and anticarcinogenic effects Generally, inhibition of mutagenicity and cancer development by polyphenolic compounds could be due to their ability to scavenge and trap potentially DNA-damaging electrophiles, free radicals and toxic metals, inhibiting enzymes that activate precarcinogens to carcinogens and inducing carcinogen-detoxifying enzymes. Reported anticarcinogenic or antimutagenic effects of both free and potato-bound chlorogenic acid include: Page 15 1. nitrites in food can react with secondary amines to form mutagenic and carcinogenic nitrosamines. Chlorogenic acid and other polyphenols are reported to block nitrosamine formation by competitively reacting with the nitrite (Kikugawa et al. 1983), 2. chlorogenic acid and several other simple phenolic acids also inactivate the mutagenicity of aflatoxin B1 (Stich & Rosin 1984), 3. chlorogenic acid can bind the carcinogen benzo[α]pyrene (Camire et al. 1995). Potato peel is better at binding this carcinogen than wheat bran, cellulose or arabinogalactan. 3.7.4 Glucose-lowering properties Thompson et al. (1983) reported that the polyphenol content of potatoes, legumes, and cereals correlated negatively with the blood glucose response (glycemic index) of normal and diabetic humans consuming them in a controlled study. The glucose-lowering effect of polyphenols may arise from their ability to inhibit amylases (which catalyse the hydrolysis of starch to glucose), phosphorylases (which are involved in starch metabolism), and proteolytic enzymes (that catalyze the hydrolysis of proteins to free amino acids in the digestive tract) and/or from their ability to direct complexation between the polyphenols and starch, preventing digestion. Another possibility is the prevention of in vivo nonenzymic browning between plasma glucose and amino groups of hemoglobin, which occurs under physiological conditions and contributes to the cause of diabetes (Friedman 1997). 3.7.5 Cholesterol-lowering effect Chlorogenic acid, and other polyphenols, also exhibit strong in vitro antioxidant activity for heart disease-related lipoproteins (LDL) (Vinson et al. 1995). Since in vivo oxidation of LDL appears to be a major cause of heart disease, it is possible that chlorogenic acid and other polyphenols may also lessen heart disease. Lazarov & Werman (1996) found that consumption of potato peel induced a lowering of cholesterol in rats. They ascribed this to its fibre content, but it is likely that the polyphenol content and other antioxidants as well as glycoalkaloids contributed to the observed hypocholesterolemia. Both tomato and potato glycoalkaloids have a strong affinity for cholesterol (Friedman et al. 1997). 3.7.6 Effect on protein nutritional quality Polyphenolic compounds and derivatives (especially tannins) bind to proteins in the gut, adversely affecting absorption of food (Friedman 1989; Oste 1989; Duffey & Stout 1996). Rat feeding studies suggest that the reduction in protein nutritional quality following consumption of polyphenols may be due to the formation of protein-polyphenol complexes (Spencer et al. 1988) and to inhibition of the digestive enzymes α-amylases and trypsin (Griffiths 1986). Polyphenols also induced an increase in lipase activity in the digestive system. Possible effects of polyphenol-rich diets on protein nutrition need further study. Page 16 3.8 Plant pigments The appearance of food is important to consumers and a major factor determining appearance is colour. Plant colour is determined by three main classes of pigments: carotenoids, chlorophylls and flavonoids (anthocyanins). Some of these compounds have come under increasing scrutiny because of their possible role in human health. Within potato cultivars there is a considerable range of colour, including white, yellow, brown, pink, red, purple, purple-blue to almost black. These colours are due mainly to carotenoids and anthocyanins. 3.8.1 Carotenoids The flesh of potato varieties is often tinged with yellow and this is mainly due to the presence of carotenoids, a class of plastid pigments. The main carotenoid constituents of potato tubers are the xanthophylls violaxanthin, lutein and lutein-5,6-epoxide, with small amounts of neoxanthin and neoxanthin-A present. β-Carotene, a common carotenoid in many other plants and also present in the aerial parts of the potato plant is absent or present in only trace amounts in the tubers (Burton 1989). There is a direct correlation between yellow flesh colour and total carotenoid content, which is a heritable characteristic. Typical “white” flesh potatoes contain 0.01-0.05 mg of carotenoids per 100 g FW while varieties with “yellow” flesh contain 0.11-0.34 mg of carotenoids per 100 g FW (Gross 1991). Data on the carotenoid content (∃-carotene equivalents) of New Zealand potatoes are give in Table 10 in Appendix I. Even the yellow-fleshed potato varieties are low compared to many kumara/sweet potato (Ipomoea batatas Lam.) cultivars where typical carotenoid levels vary between 0.10 and 7.5 mg per 100 g FW, and some very dark orange cultivars even have up to 20 mg of carotenoids per 100 g FW. It is thought that the tendency for a high carotenoid content is determined by a single dominant gene, although there are modifying genes (Brown et al. 1993). 3.8.2 Anthocyanins The other major group of pigments which occur in some potato cultivars is the anthocyanins. These have been studied comprehensively by Lewis (1996). The anthocyanins are vacuolar pigments which give rise to the red to purple/back colour of tuber skin and sometimes flesh. The degree of tuber flesh pigmentation can vary considerably from just a slight pigmentation of the vascular ring to a complete pigmentation of the entire tuber. Anthocyanins are important antioxidants. Red cultivars of some vegetables often have stronger antioxidant activity than their non-coloured relatives. Studies with Red Desiree (red-skinned) potatoes have shown them to have higher antioxidant activity than Rua (Lister & Podivinsky 1998). However, since the skin only accounts for a relatively low percentage of the total weight there were not huge differences (approximately 20% higher), unlike lettuce where a red cultivar had activity 13 times that of a green cultivar. It is possible that a red-fleshed potato would have much higher antioxidant activity than Page 17 those with only red skin. However, there have been no studies of these aspects to date. 3.8.3 Others Greening in potatoes is due to chlorophyll formation as a result of exposure to light. It is usually accompanied by the production of the toxic glycoalkaloids (see Section 3.3.4). Melanin formation as a result of injury may give rise to grey or black discolorations, which are also undesirable. 3.9 Nutrient losses/changes during cooking Some of the effects of cooking on specific nutrients have been discussed in the previous sections. However, other issues arise. Recent consumer interest in 'healthy eating' has raised awareness to limit the consumption of fat and fatty foods. An often asked question is, “What are the relative nutritional advantages and disadvantages of consuming fried foods?” Are all fried foods bad for you? The nutrient losses and gains during frying have been reviewed by Fillion & Henry (1998). Frying generally has little or no impact on the protein or mineral content of fried food, whereas the dietary fibre content of potatoes is altered after frying due to the formation of resistant starch. The high temperature and short transit time of the frying process cause less loss of heat labile vitamins than other types of cooking. Frying also avoids loss of water-soluble vitamins through leaching into the cooking water. For example, vitamin C concentrations of French fried potatoes can sometimes be as high as in raw potatoes, and thiamine is well retained in fried potato products as well as in fried pork meat. The nutritive value of the frying media is also important to take into consideration as it can contain beneficial compounds such as vitamin E. Although some unsaturated fatty acids and antioxidant vitamins are lost due to oxidation, fried foods can be a good source of vitamin E. Of course this depends on the type of oil they are cooked in and only some of the New Zealand fries had higher vitamin E content (see Table 10 in Appendix I). It is true that some fat is inevitably taken up by the food being fried, contributing to an increased energy density. However, this also results in highly palatable foods with a high nutritional content. It has been concluded that fried foods certainly have a place in our diets in moderation (Fillion & Henry 1998). Attempts to reduce the oil content of the potatoes while retaining the other benefits of frying may be desirable, and it is preferable to use an oil with a good vitamin E content. Recent studies have indicated that cooking increases the bioavailability of some nutrients, especially carotenoids. This may be due to the breakdown and softening of cell walls, making the cell contents more readily available for absorption. Since potatoes are always cooked, some components may be more bioavailable from potatoes than from vegetables eaten raw. Page 18 4 The health benefits of potatoes There are numerous misconceptions about the nutritional value of the potato. It is often believed that the potato is a high-energy food that provides little else in the way of nutrients. This is partly due to the high satiety value of the potato and its description in food composition tables and numerous publications as a “starchy tuber”. It is sometimes regarded that potato should not be classed as a vegetable in terms of contributing to the 5 plus fruit and vegetables a day recommendations. However, in view of current recommendations that the intake of fats in the diet should not exceed about 30% of energy, that free sugar intakes should be reduced, that about 55% or more of our energy should be derived from carbohydrate, that energy intakes overall should be reduced, potatoes are an excellent base to the diet. A daily intake of 150-300 g of potatoes provides 4-8% of the calories required by a human. Potatoes are a considerably richer source of nutrients than of energy. Many of the health benefits of the components of potatoes are discussed in the previous section. The contribution potatoes make to daily energy and nutrient intakes is shown in Table 7. The benefits of potatoes can be summarized as: 1. they are virtually free of fat, although they are quite easily turned into fatty foods, 2. they contain very little soluble sugar, 3. nearly all of the carbohydrate in them is in the form of hydrated starch. Starch contributes about 17 kJ/g energy compared with 37 kJ/g for fat, and starch is hydrated because in cooked potato kJ per gram are even lower. Therefore, the volume that potato occupies per kilojoule is large, that is, the energy density of potato is low and it is able to displace fatty materials from the diet, thus reducing energy intake without a feeling of emptiness. Potato, therefore, has a role to play in managing the obesity that is contributing to the current burden of disorders such as diabetes and heart disease, 4. although the starch in potatoes is readily digested, as indicated by the high glycaemic index values, the high water content of potatoes means that on a weight for weight basis they have a relatively low impact on blood glucose - about one-third that of bread (Relative glycaemic potency of bread = 30, of potato = 10) (Monro 1999), 5. potatoes should not be relied on as a source of dietary fibre as they contain little of it, and it is easily fermented. This does, however, have some benefits because it means that people with extremely high energy intakes, such as athletes, do not become overburdened with the non-digestible food residues, and furthermore the fermentability of potato fibre means that it can contribute extra energy as short-chain fatty acids, 6. they are a good source of several water soluble vitamins. They may make a significant contribution to the daily intake of vitamin C Page 19 (approximately 30% of the recommended daily intake). They also contain important B group vitamins, especially thiamin, and folic acid, 7. potatoes are rich in minerals, particularly potassium, but are low in sodium which is a desirable balance for a healthy diet. Other important minerals include iron, 8. potatoes contain high quality protein, although they are deficient in the essential amino acid methionine, Table 7: Potato consumption and contribution to energy and nutrient intake (% recommended daily intake) (all data apart from New Zealand sourced from Lisiska & Leszcyski 1989). Country New Zealand1 Great Britain USA Consumption (g/day) 180 186 150 200 Energy Protein Carbohydrates Dietary fibre Fat Vitamin C Thiamin (B1) Riboflavin (B2) Niacin (PP) Pantothenic acid Vitamin B6 Folic acid Vitamin A Potassium Phosphorus Calcium Magnesium Iron Copper Zinc Iodine Fluorine 6.3 8.0 0.03 46.4 13.0 5.2 8.9 12.5 13.7 0.1 27.7 7.3 1.8 10.9 12.7 4.1 - 4.4 3.9 8.7 0.0 24.1 10.1 3.0 8.6 0.0 7.7 - 4.0 4.7 56.6 8.7 3.6 12.1 16.4 4.9 7.3 7.8 5.2 16.9 3.9 15.2 - 5.5-6.6 -2 16.7 0.3 48.6 18.3 6.7 16.3 10.0 26.3 3.5 22.0-29.3 2.4 16.7 8.9 7.5-15.0 2.0 Specification West Germany 1 Figures are based on nutrient composition data from the NZ Food Composition Database (average of potatoes cooked by different methods) and NZ RDI values for adults (average of male and female requirements and of average weight). 2 No data available. 3 Unless eaten in a fried form which will make some contribution. 9. phenolics are present in fairly low quantities in typically consumed potato cultivars but still may have an important role in health because of the quantities of potatoes consumed compared to many other vegetables. Not enough is known to be sure of their benefits, Page 20 10. 5 glycoalkaloids are an undesirable component in potatoes but the risk of toxic levels causing harm to human health is minimal. The future Much of the focus on potato breeding and development in the past has been focused on increasing yields and developing pest and disease resistance. In addition to generating new traits that enable the plant to grow better (input traits), which are useful to farmers, plant breeding and genetic engineering technology can also generate plants with improved nutritional features (output traits). Although the nutritional value of the potato is fairly high there is still plenty of room for improvement. Most other vegetables (e.g. broccoli, tomatoes, carrots) are regarded as having some health benefits that outweigh that of potatoes. However, it is much easier to get a child to eat a potato than say a piece of broccoli. Therefore the question arises, “Can the health benefits of broccoli be incorporated into potatoes?” Some manipulation of nutritional characteristics is possible through traditional plant breeding and cultural practices. However, genetic engineering presents greater opportunities. Genetic modification of potatoes is relatively easy because they are amenable to transformation using Agrobacterium tumefaciens, and plantlets are readily regenerated and clonally propagated. 5.1 Feeding the world in the twenty-first century One of the major problems facing mankind is the need to feed an increasing population, particularly in developing countries. As well as gross undernourishment, lack of protein, vitamins, minerals and other micronutrients in the diet is also widespread. Although these problems are mainly evident in developing countries, the health of many New Zealanders and people in developed countries around the globe is severely compromised by diseases that afflict affluent societies (e.g. heart disease, cancer). These disease have increased significantly over the last few decades and are often associated with high levels of obesity and blood cholesterol. Ironically, large sections of these populations are "malnourished", not in the classical sense of having insufficient food but in terms of not eating a balanced diet, with food of good nutritional quality, to aid in the prevention of disease; the World Health Organization has reported that more than 30% of non-communicable disease can be prevented by diet. As a response to this there has been a huge upsurge in the supplement industry. However, these supplements do not always deliver the same health benefits as eating foods of high nutritional quality. Thus, it is important to find ways to improve the nutritional quality of food that is easily distributed or grown in developing countries and/or regularly consumed by the majority of the population. Page 21 5.1.1 Protein quality improvement When diets are high in carbohydrates and low in protein over a long period, essential amino acid deficiency results. Many foods, especially those of plant origin, such as potatoes, have low levels of amino acids which limit their nutritive value. Inadequacies in nutritional value of protein can be solved in at least three ways: (a) combining protein sources to create mixtures with an adequate amino acid balance, (b) fortification of the low quality proteins with essential amino acids, and (c) developing high quality plant proteins by breeding or molecular biology techniques. As discussed in Section 3.3.1, although potatoes contain high quality protein they lack essential amino acids. Small changes may be possible by selection. However, to make significant differences strategies for manipulation by genetic engineering are needed. Tu et al. (1994) attempted to incorporate the Brazil nut 2S gene into potato. This gene was expected to express a protein with elevated methionine and cysteine, but the expression levels observed in tubers of engineered plants were insufficient to increase their methionine content. With the knowledge that amino acid synthesis in higher plants is controlled by feedback inhibition, Widholm (1977) used the appropriate analogue for an amino acid to select cell lines with relaxed feedback control of amino acid biosynthesis and to overproduce the protein. For example, a carrot cell line accumulated 10 times the normal level of free methionine. A similar protocol was used by Langille et al. (1998) to select protoclones of Russet Burbank potatoes that had up to three times the free methionine content of the control. It has not been determined whether this trait remains stable in subsequent generations. Genes that code for proteins with a high content of the essential amino acids found to be most deficient in plant-derived proteins have been constructed and expressed in bacteria (Jaynes et al. 1985). Since then, research has been directed towards developing potato plants with these synthetic proteins. There have been some problems with expression levels of the new proteins (Destefano-Beltran et al. 1991). More recently a US patent has been filed which describes the insertion of a gene responsible for producing a protein in potatoes and rice that is relatively high in essential amino acids (Jaynes & Derrick 1998). Theoretically, proteins could be specifically designed to supplement any desired animal feed or human food. Insertion of lysine at frequent intervals in synthetic proteins provides sites for proteolytic attack by Trypsin (a digestive enzyme), increasing the bioavailability of the supplemental protein (Destefano-Beltran et al. 1991). 5.1.2 Provitamin A Vitamin A deficiency is one of the major vitamin deficiencies in developing countries. A lack of vitamin A can lead to eye damage and even blindness and also weaken the protective barriers to infection put up by the skin, the mucous membranes and the immune system (Somer & West 1966). One of the most exciting recent developments has been the introduction of genes into rice that result in the production of the vitamin A precursor β-carotene in the grain (Conway & Toenniessen 1999). β-carotene is a pigment required for Page 22 photosynthesis and is synthesized in the green tissues of all plants, including rice and potatoes. However, it is not always present in non-photosynthetic tissue such as seeds and tubers. No rice mutants have been found that produce β-carotene in the grain so traditional breeding is not an option. Therefore, genes encoding for β-carotene synthesis were incorporated into rice by genetic engineering. The grain has a light golden-yellow colour and contains sufficient β-carotene to meet human vitamin A requirements. Although some potato cultivars are yellow they do not produce much β-carotene and thus have no or low provitamin A content. Strategy will be different that that used for rice need to alter composition of carotenoids. Problems with consumer acceptance of yellow potato however those parts of the world where food is a major doesn’t matter. 5.1.3 Disease and pest resistance To date the large focus on genetic engineering of potatoes has been to develop cultivars that are resistant to pests and diseases. The development of such plants will reduce pesticide usage, benefiting human health. The volume of chemicals remaining on potatoes at consumption is likely to be small and in most cases insignificant in terms of human health. However, there are huge health benefits for growers who do not to have to spray chemicals. 5.1.4 Minerals Levander (1990) has discussed the possibilities for developing fruit and vegetables with altered mineral composition. Fruits and vegetables contribute relatively little selenium or molybdenum to the total dietary intake. Selenium may play a protective role against certain human cancers. Should it prove desirable to increase dietary intake of selenium, vegetables could be an important component of such a strategy. A variety of vegetables (potato, tomato, carrots, cabbage, and onion) grown in seleniferous soils may contain two to three orders of magnitude more selenium than those grown in nonseleniferous soils. New Zealand tends to have low selenium levels in its produce because of the low levels in its soils. Currently, potatoes are a very poor source of selenium; they contain on average 0.63 μg/100 g. A 180 g serving of potatoes only contributes about 1.5% of the recommended daily intake of selenium. The desirability of increasing selenium in the food supply in general is still uncertain. Under some experimental conditions, elevated intakes of selenium actually increased rather than decreased the incidence of chemically induced cancer in laboratory animals. Thus, in certain situations selenium may not only be ineffective, but also harmful. However, the levels that have adverse effects are far higher than those that may be achieved from a high intake of fruit and vegetables. Public health authorities in various countries must exercise caution when deciding whether or not to add selenium to their national food supplies. Nutrient elements in other crops have been manipulated. For example, genes have been added to rice to increase the grain’s available iron content (Conway & Toenniessen 1999). Such approaches could be applied to Page 23 potatoes. Potatoes are a significant source of iron in the New Zealand diet. However, some females do not have adequate iron intake so there is room for improvement. On the other hand, iron plays a role in after-cooking darkening so it may not be desirable to increase the level. 5.2 Delivery of additional health benefits Functional foods are gaining more and more interest from health scientists. Potatoes may be a good way to deliver health benefits because of their levels of consumption in most people’s diets. 5.2.1 Increased levels of phenolics and other antioxidants There have been many promotions to increase fruit and vegetables in the diet such as the 5+ a day promotion. More recently evidence suggests that people should be eating at least eight servings a day, especially those with a predisposition to diseases such as heart disease and cancer. One of the key components of fruit and vegetables that may help prevent diseases are antioxidants (e.g. vitamins C and E, phenolics, carotenoids). Although potatoes have reasonable levels of vitamin C they have relatively low levels of phenolics and white-fleshed varieties have only traces of carotenoids. Thus, the antioxidant activity of potatoes is much lower than many other vegetables. By increasing the levels of antioxidants in commonly eaten vegetables it may be possible to obtain the health benefits from consuming a more moderate number of servings which may be achieved by a greater proportion of the population. Some increase in antioxidant levels is achievable through traditional breeding and using particular cultural practices. In addition there are opportunities for the genetic manipulation of antioxidants in plant foods and some of these have been discussed by Mullineaux & Creissen (1996). There has been some traditional breeding of potatoes for increased vitamin C content. Through an understanding of the biosynthesis of ascorbic acid it may also be possible to manipulate the levels through genetic engineering. While the vitamin C content of potatoes is quite reasonable the levels required to have other beneficial effects on health are likely to be much greater than those required to avoid vitamin deficiency. High doses probably should be avoided although any excess of water-soluble vitamins is simply excreted in urine. Intakes of vitamin C of 1 gram per day have had no adverse effects. However, it is now thought that the best health benefits, particularly those from antioxidants, are gained from consuming a range of different antioxidants rather than large doses of a single compound. Therefore, a better approach may be to manipulate the levels of other natural antioxidants. Another way to raise the antioxidant activity of potatoes is to increase the levels of phenolics. Potatoes with a high phenolic content would have to be acceptable to consumers with respect to blackspot, after-cooking darkening and sensory qualities. The consumer may have to choose between perceived undesirable appearance and real beneficial health effects. Another important consideration is that some phenolics have a strong taste, mainly bitter, so Page 24 increasing their levels may have undesirable effects on flavour. Thus, these factors will need to be carefully balanced. The following is a list of some aspects of potato phenolics that need investigation. 1. Define the relative health promoting properties of the various polyphenols present in potatoes and determine if there are synergistic effects. 2. Examine the antioxidant activity of coloured potato cultivars to see if these have increased antioxidant activity over non-coloured varieties. 3. Enhance the content of the most potent antioxidative polyphenols in potatoes either by plant breeding or molecular biology techniques. It may be particularly valuable to increase the levels in the flesh so that peeling does not adversely affect the health benefits. It may also be advantageous to increase the range of compounds present rather than just increase the levels of chlorogenic acid. 4. Define the effects of commercial and home food processing on polyphenols in potatoes. 5. Investigate possibilities for enhancing chlorogenic acid content, and other phenolics, by postharvest exposure to light and other stress conditions (although care would be required to avoid glycoalkaloid accumulation). 6. Develop new, inexpensive, nutritious and health-promoting highchlorogenic acid/phenolic potato tuber, peel and leaf food formulations. The value of leaves as a possible food source has received some attention. Edible leaf protein isolates have been prepared previously and generally major efforts have been made to remove the pigments (Bickoff et al. 1973; Pirie 1973; Friedman 1996a). Since leaves have a high polyphenol content and since chlorophyll binds strongly to carcinogenic heterocyclic amines (Friedman 1996b) it may be worthwhile to retain both of these leaf constituents in the preparation of isolates. 7. Carry out animal and human feeding studies with high phenolic potatoes to assess whether beneficial effects of chlorogenic acid, and other phenolics, observed in vitro are confirmed in vivo. The other group of antioxidants that could be manipulated in potatoes are the carotenoids. The manipulation of carotenoids has already been discussed in Section 5.1.2 regarding provitamin A activity. The same principal could be used to increase total carotenoids (not necessarily just ∃-carotene) in potatoes for their antioxidant properties. In many European countries yellow potatoes are quite acceptable and in fact often preferred, but in other countries, such as New Zealand, a white potato is usually the norm. Page 25 5.2.2 Recombinant proteins/vaccines A large number of recombinant proteins are being developed for pharmaceutical purposes and will have to be produced on a large scale for clinical studies and pharmaceutical applications. Protein farming in transgenic plants offers great flexibility at very low primary cost as expensive technical cell culture facilities are not required (Artsaenko et al. 1998). Furthermore, contamination of the product by mammalian viruses or bacterial endotoxins is excluded. Therefore, transgenic plant-based production systems are a competitive alternative to bacterial or mammalian cell culture-based systems (Goddijn & Pen 1995). Potato tubers have great potential in biofarming because those storage organs can be used to accumulate large amounts of protein. Vaccination against infectious diseases has revolutionalised human health. Diseases such as smallpox have been eliminated and others such as polio have almost disappeared. There has been a significant increase in our understanding of immunology and in the development of new technologies, increasing hope that new vaccines will be developed for other diseases. Recent vaccine developments have focused on finding ways of delivering various types of vaccines. An approach that has great appeal in relation to the potential ease of both production and delivery is using transgenic edible plants to make vaccines. They may provide a more convenient and practical means of implementing universal vaccination programmes throughout the world. For example, genes encoding antigens have been cloned into potato plants. The potato makes the protein. When animals and more recently humans eat the potato they develop immune responses (Liu 1999). The drawback of using potatoes is that they are usually eaten cooked, and this may denature a protein antigen. In addition to having a role in combating disease, plant-derived vaccines may have potential as a means of contraception (Smith et al. 1997). The public acceptability of plant vaccines has been considered by Danner (1997). Some examples currently being developed using potatoes either as a test system or as a final delivery system are noted below. ϒ A transgenic plant was shown to be capable of synthesizing an immunogenic form of the diabetes-associated autoantigen, glutamic acid decarboxylase. Given as a dietary supplement, the transgenic plant inhibited the development of diabetes in the non-obese diabetic mouse (Ma et al. 1997). ϒ Potato has been used to produce an edible vaccine protecting against enterotoxigenic E. coli (Mason et al. 1998). The vaccine has been expressed in leaves and tubers. Trials with mice have shown some degree of protection against E. coli. However, the rats were fed raw potato which would not be possible in human trials. A trial with a small group of humans showed antibody production (Tacket 1998). ϒ Transgenic potatoes have been developed in an attempt to protect against chlorea (Arakawa et al. 1998). Some degree of protection has been demonstrated in mice. Page 26 ϒ 5.2.3 Research is currently underway to develop transgenic potato plants for use as an oral vaccine against cryptosporidiosis, an important cause of diarrhoea worldwide (Hudspeth 1997). Modified carbohydrate One way to improve the nutritional properties of foods may be to modify the starch component, often included to stabilize the food product. High value carbohydrates produced in plants include bacterial cyclodextrins that have been produced in transgenic potato tubers (Oakes et al. 1991). There are various applications for cyclodextrins, including pharmaceutical delivery systems, flavour and odour enhancement and the removal of undesired compounds (such as caffeine) from foods. Modified potato starches have been tested for their dietary effects (Raben et al. 1997). Two chemically modified starches—a 1-2% acetylated potato starch and a starch enriched with 2% beta-cyclodextrin—and a native, unmodified potato starch (control) have been investigated with regard to 6 hour energy expenditure, substrate metabolism, hormone concentrations, and subjective appetite sensations. A flattening of the glucose curve, a lower insulin and gastric inhibitory polypeptide response, and higher fullness ratings were observed after the meal with the beta-cyclodextrin starch. Satiety ratings were higher after both meals with modified starch than after the meal with the control starch. It was concluded that a minor modification of native potato starch improved the glycemia, insulinemic, and satiating properties of a meal. Slower gastricemptying rate or delayed intestinal absorption of the modified starch may explain the observed differences. 5.2.4 Other A gene encoding the human milk protein beta-casein has been introduced into potato, and expression of human milk beta-casein was demonstrated in both leaf and tuber tissues (Chong et al. 1997). These findings open the way for reconstitution of human milk in edible plants to replace bovine milk in baby foods for general improvement of infant nutrition and to prevent gastric and intestinal diseases in children. The gene for human epidermal growth factor (hEGF) was chemically synthesized and has been successfully expressed in transgenic potato (Salmanian et al. 1996). 5.3 Removal of antinutritional compounds and undesirable chemicals 5.3.1 Glycoalkaloids Potatoes contain glycoalkaloids which have detrimental effects on human health. Care must be taken when producing new varieties of potatoes so that the level of glycoalkaloids does not rise to an unsafe level or that new, more toxic glycoalkaloids are not introduced into the commercialised germplasm (Friedman & McDonald 1997). Strategies must be developed to minimize postharvest glycoalkaloid production. These may include: Page 27 1. determining relative susceptibilities to greening and mechanical damage of current and new cultivars, 2. evaluating food-compatible enzyme inhibitors, such as citric acid, and substrate inhibitors for their ability to inhibit enzymes catalysing glycoalkaloid biosynthesis, 3. evaluating films with built-in chromophores that absorb light and protect potatoes against greening, browning and spoilage, 4. investigating the effects of size and maturity on glycoalkaloid levels and measuring glycoalkaloid changes during sampling and handling. In addition, studies are required to reduce preharvest glycoalkaloid formation by suppressing genes governing their biosynthesis. This will provide a variety of benefits extending from the growing, processing, shipping and marketing to the consumption of potatoes and potato products. Reduction of toxicant levels in selected varieties will allow the introduction of new potato cultivars that cannot currently be released due to their higher than acceptable levels of glycoalkaloids. This would enhance the value of potatoes as a high quality food. Genetic engineering may allow the creation of new cultivars without altering other desirable characteristics. Approaches may use antisense RNA and related methods to develop cultivars with low levels of glycoalkaloids while maintaining acceptable resistance to phytopathogens (Friedman & McDonald 1997). The gene encoding solanisine glucosyl transferase is one target. At present it is unclear how lowering glycoalkaloid production would affect the concentrations of other compounds, such as phytoalexins, which may have major consequences for pest resistance. 5.3.2 Chemical residues In addition to possible herbicide and pesticide residues, potato peels have been shown to harbour significant amounts of chemical residues that may be mildly toxic. In the USA chemical sprout inhibitors have been found in significant levels in potato peels, even after cooking (Lang 1992). Sprout inhibitors are important chemicals that help prevent shrinkage, blackening, nutrient loss, and susceptibility to bruising, and may even reduce the accumulation of some natural toxic chemicals that accompany sprouting (see section on glycoalkoids). Peeling potatoes prior to cooking removes most of the problems associated with toxic substances in the peel. However, the peel contains many beneficial components. Strategies for natural prevention of sprouting by genetic engineering or other means are desirable. Prevention of sprouting may also help prevent the accumulation of glycoalkaloids during storage. 5.4 Browning prevention There are a number of strategies to minimise or prevent browning and these require continued study. One approach is the breeding of potatoes with an initial low chlorogenic acid content to minimize both content and rate of increase, e.g. to minimize after-cooking darkening and/or enzymic browning during food processing. However, since chlorogenic acid is reported to have Page 28 beneficial effects on health (as described in Section 3.7), it may be desirable to breed for high chlorogenic acid varieties. A construct expressing a high tyrosine protein has been introduced to potato to act as a tyrosine-sink and decrease the amount of free tyrosine which contributes to browning reactions (Vayda & Belknap 1992). Other efforts include employing antisense constructs of potato polyphenol oxidase genes (Bachem et al. 1994). Understanding the chemical, nutritional and toxicological consequences of browning reactions and related transformations can lead to better and safer foods and feeds and improved human health. 5.5 Other new and improved potato products Over the last few years there has been an increase in the number of potato products appearing on the market. With the increasing trend to convenience and processed products there is a need to ensure that nutritional value is not lost during processing and storage of these foods. 5.5.1 “Healthier” fried products There is a high consumption of potatoes in the form of chips, French fries, etc. Yet with the increasing levels of obesity there is a need to reduce consumption of fatty foods by at-risk sections of the population. Apart from cutting fried foods out of the diet there are various strategies for achieving a lower fat content, including altering the cooking media and altering the composition of the potato. One advancement has been the development of olestra, a non-absorbable fat substitute. A recent study using this material showed that substituting fat-free (olestra-containing) potato chips for regularfat chips can help reduce fat and energy intakes in short-term (Miller et al. 1998). However, the use of olestra is being debated and requires further investigation. Olestra is reported not to be toxic, carcinogenic, genotoxic, or teratogenic, and is neither absorbed nor metabolised by the body, but may be associated with gastrointestinal tract symptoms such as cramping or loose stools (Prince & Welschenbach 1998). In addition, olestra affects the absorption of fat-soluble vitamins but does not affect the absorption of watersoluble nutrients. The petitioner's studies concluded that when olestra was consumed with foods containing vitamins A, D, E, or K, the fat substitute could have an effect on the absorption of these nutrients. Other strategies for reducing fat intake include modification of starch genes to produce potatoes with a higher dry matter (Anon. 1992). This results in less water for oil to replace in processing and thus results in a “healthier” fried food. Increased solids contents through genetic modification have already been reported (Fraley 1991; Fraley et al. 1991) and there is potential for further increases. 5.5.2 Novelty coloured potato products The array of coloured potatoes that are available, or could be engineered, presents opportunities for the production of novelty coloured potato products. These may be red, blue or purple due to anthocyanins or yellow/orange from carotenoids. In addition to the colour such products may have health benefits Page 29 due to the antioxidant activity and other modes of action of the anthocyanins and carotenoids. Details of anthocyanins and carotenoids and possible manipulation of these attributes have been discussed in earlier sections (3.7, 3.8 and 5.2.1). If coloured potatoes are to be used it is important that they retain their colour during cooking and that the final product is highly coloured and retains its nutritional benefits. During normal cooking (boiling, steaming or crisping) there was no thermal degradation of anthocyanins and intensely coloured products were obtained (Lewis 1996). The only significant loss of anthocyanin occurred by leaching from cut surfaces when potatoes were boiled. Therefore, coloured tubers should be cooked whole and cut up after cooking. The effects of processing on the antioxidant activity of potatoes have not yet been evaluated and will need to be if products are to be developed. 5.6 Utilizing potato waste New opportunities for utilizing potato waste have been explored by Heyes et al. (1997) and so these are not discussed in any detail here. Uses include: ϒ stockfeed: increasing importance particularly if some of the nutritional qualities are manipulated as discussed above, ϒ specialty potato products (e.g. potato mash, flakes and granules for use in potato novelties or extruded foodstuffs), ϒ potato flour improves the nutritional value of bakery products, ϒ extraction of potato protein, ϒ pigment extraction: natural pigment to replace artificial and also added health benefits, ϒ extraction of nutraceuticals: extraction of potato starch–has potential as a fat replacer in foods such as sausages making them a healthier option. Another usage of potato peels in particular could be as a source of a natural antioxidant preservative with wide food applications. Potato peel extracts have also been shown to have bacteriocidal and bacteriostatic effects (Rodriguez de Sotillo et al. 1998). 6 Recommendations/future research This review identifies a number of potential areas for future research. Genetic engineering offers huge potential for manipulating many different aspects of nutritional quality and introducing additional health benefits. In light of the current debate, and a possible moratorium on genetic engineering, these will need careful consideration. However, the public may be more likely to accept a genetically engineered vegetable when they can see a personal benefit. Despite this there is still considerable scope for improvements through traditional plant breeding and/or changes in agronomic and postharvest handling procedures. Page 30 Future projects could include: 7 ♣ developing strategies for improving the antioxidant activity of potatoes. There are a number of ways to achieve this but first different varieties of potatoes should be screened and particularly those with coloured flesh. Levels of phenolics could be increased by traditional breeding and genetic engineering, ♣ increasing the provitamin A content of potatoes. Some improvements may be possible through traditional breeding and selection of cultivars, but to achieve significant increases genetic modification would be required. Marketing surveys will be required to determine if New Zealand consumers would accept a yellow potato, ♣ developing potatoes with high anthocyanin and/or carotenoid contents creates opportunities for the development of novelty coloured potato products (e.g. extrusion products, crisps), which may have the added health benefits of the compounds they contain. If such products are to be developed then the effects of processing must be carefully investigated to ensure the health benefits are not lost, ♣ producing higher dry matter potatoes for processing to reduce the fat content of fried products, ♣ reducing glycoalkaloid content of potatoes. In addition it is important to monitor the glycoalkaloid content of new cultivars as there is a risk of increased glycoalkaloid biosynthesis, although this risk is very small, ♣ as new cultivars are developed with additional health benefits there will be a need to re-evaluate the potential of potato wastes to better utilize the key components either for improvement of human or animal health, ♣ evaluating the possibilities of using genetic engineering to produce vaccines and high value pharmaceuticals from potatoes. Since we have very strong programmes for genetic modification of potatoes these could easily incorporate identified opportunities, ϒ developing new potato cultivars with clearly defined nutritional and health benefits opens up potential for the development of tailored products for specific sectors of the population (e.g. diabetics, the elderly, athletes). Market opportunities and potential product development to meet changing New Zealand and world demands (e.g. as a result of the aging population) will need to be considered. Acknowledgements The authors would like to thank Dr Karen Silvers and Emmeline Taptiklis for the nutritional information, and Dr Tony Conner for information and literature on genetic engineering opportunities for potatoes. Page 31 8 References Adams, J.B. 1994: Green colour development in potato cooking water. Food Chemistry 49: 295-298. Al-Saikhan, M.S.; Howard, L.R.; Miller, J.C. 1995: Antioxidant activity and total phenolics in different genotypes of potato (Solanum tuberosum L.). Journal of Food Science 60: 341-343. Anon. 1992: The starch genes. Spudman 9/92: 13-15. Arakawa, T.; Chong, D.K.X.; Langridge, W.H.R. 1998: Efficacy of a food plant-based oral cholera toxin B subunit vaccine. Nature Biotechnology 16: 292-297. Artsaenko, O.; Kettig, B.; Fielder, U.; Conrad, U.; During, K. 1998: Potato tubers as a biofactory for recombinant antibodies. Molecular Breeding 4: 313319. Bachem, C.W.B.; Speckmann, G.-J.; van der Linde, P.C.G.; Verheggen, F.T.M.; Hunt, M.D.; Steffens, J.C.; Zabeau, M. 1994: Antisense expression of polyphenol oxidase genes inhibits enzymatic browning in potato tubers. Biotechnology 12: 1101-1105. Bickoff, E.M.; Booth, A.N.; de Fremery, D.; Edwards, R.H.; Knuckles, B.E.; Miller, R.E.; Saunders, R.M.; Kohler, G.O. 1973: Nutritional evaluation of alfalfa leaf protein concentrate. In: Protein Nutritional Quality of Foods and Feeds, Vol 2. Friedman, M., Ed. Dekker, New York. Pp. 319-340. Bird, A.R. 1999: Prebiotics: A role for dietary fibre and resistant starch? Asia Pacific Journal of Clinical Nutrition 8(Suppl.): S32-S36. Brown, C.R.; Edwards, C.G.; Yang, C.-P.; Dean, B.B. 1993: Orange flesh trait in potato: inheritance and carotenoid content. Journal of the American Society for Horticultural Science 118: 145-150. Burlingame, B.A.; Milligan, G.C.; Spriggs, T.W.; Athar, N. 1997: The Concise New Zealand Food Composition Tables 3rd Edition. New Zealand Institute for Crop & Food Research Ltd. Burton, W.G. 1989: The potato, 3 rd Edition, Longman Scientific and Technical, Harlow, Essex, UK. Bushway, R.J.; Bureau, J.L.; McGann, D.F. 1983: ∀-Chaconine and ∀solanine content of potato peels and potato peel products. Journal of Food Science 48: 84-86. Camire, M.E.; Zhao, J.; Dougherty, M.P.; Bushway, R.J. 1995: In vitro binding of benzo[α]pyrene by extruded potato peel. Journal of Agricultural and Food Chemistry 43: 970-973. Chong, D.K.X.; Roberts, W.; Arakawa, T.; Illes, K.; Bagi, G.; Slattery, C.W.; Langridge, W.H.R. 1997: Expression of the human milk protein beta-casein in transgenic potato plants. Transgenic Research 6: 289-296. Page 32 Conway, G.; Toenniessen, G. 1999: Feeding the world in the twenty-first century. Nature 402 (Supp): C55-C58. Danner, K. 1997: Acceptability of bio-engineered vaccines. Comparative Immunology and Microbiology of Infective Diseases 20: 3-12. Destefano-Beltran, L.; Nagpala, P.G.; Kim, J.; Dodds, J.H.; Jaynes, J.M. 1991: Use of synthetic genes to enhance nutritional quality and disease resistance in plants: application to potato. In: Molecular Methods for Potato Improvement. Report of the Planning Conference on “Application of Molecular Techniques to Potato Germplasm Enhancement”. Pp. 49-59. Duffey, S.S.; Stout, M.J. 1996: Antinutritive and toxic component of plant defense against insects. Archives of Insect Biochemistry 32: 3-37. Englyst, H.N.; Cummings, J.H. 1987: Digestion of polysaccharides of potato in the small intestine of man. American Journal of Clinical Nutrition 45: 423431. Fillion, L.; Henry, C.J. 1998: Nutrient losses and gains during frying: a review. International Journal of Food Science and Nutrition 49: 157-168. Fraley, R.T. 1991: Commercialization of genetically modified plants: progress towards the marketplace. Abstracts of the Third International Congress of the International Society for Plant Molecular Biology 3, Tuscon, Arizona, USA. Fraley, R.T.; Perlak, F.J.; Fischoff, D.A.; Tumer, N.; Stark, D.; Barry, G.; Kishore, G. 1991: Improving potato processing and pest control through gene transfer. Abstracts of the 2nd International Potato Molecular Biology Symposium, p. 28. St Andrews, Scotland. Friedman, M. 1989: Absorption and utilization of amino acids. CRC Press, Boca Raton, Florida. Friedman, M. 1992a: Dietary impact of food processing. Annual Review of Nutrition 12: 119-137. Friedman, M. 1992b: Composition and safety evaluation of potato berries, potato and tomato seeds, potatoes, and potato alkaloids. ACS Symposium Series 484: 429-462. Friedman, M. 1996a: Nutritional value of proteins from different food sources. A review. Journal of Agricultural and Food Chemistry 44: 6-29. Friedman, M. 1996b: Food browning and its prevention: an overview. Journal of Agricultural and Food Chemistry 44: 631-653. Friedman, M. 1997: Chemistry, biochemistry, and dietary role of potato polyphenols. A review. Journal of Agricultural and Food Chemistry 45: 15231540. Friedman, M.; Dao, L. 1992: Distribution of glycoalkaloids in potato plants and commercial potato products. Journal of Agricultural and Food Chemistry 40: 419-423. Page 33 Friedman, M.; Fitch, T.E.; Levin, C.E.; Yokoyama, W.H. 1997: Reduction of dietary cholesterol in hamsters fed tomatine. Presented at the National Meeting of the American Chemical Society. Abstract AGFD 81. Friedman, M.; McDonald, G.M. 1997: Potato glycoalkaloids: chemistry, analysis, safety, and plant physiology. Critical Reviews in Plant Sciences 16: 55-132. Garcia-Alonso, A.; Goni, I. 2000: Effect of processing on potato starch: in vitro availability and glycaemic index. Nahrung-Food 44: 19-22. Goddijn, O.J.M., Pen, J. 1995: Plants as bioreactors. Trends in Biotechnology 13: 379-387. Griffiths, D.W. 1986: The inhibition of digestive enzymes by polyphenolic compounds. In: Nutritional and Toxicological Significance of Enzyme Inhibitors in Foods, Friedman, M., Ed. Plenum, New York. Pp. 509-515. Gross, J. 1991: Pigments in vegetables. Van Nostrand Reinhold, New York. Pp. 208-225. Harris, P.H. 1999: Dietary fibre and the prevention of colorectal cancer. Third Australasian Dietary Fibre Workshop, 21-23 March, 1999. Hasegawa, D.; Johnson, R.M.; Gould, W.A. 1966: Effect of cold storage on chlorogenic acid content of potatoes. Journal of Agricultural and Food Chemistry 14: 165-169. Heyes, J; Genet, R.; Brash, D.; Doens, C. 1997: Opportunities for alternate uses of potatoes in New Zealand. CropInfo Confidential Report No. 450. Prepared for NZ Vegetable & Potato Growers Federation Inc. Horton, D.; Sawyer, R.L. 1985: The potato as a world food crop, with special reference to developing areas. In: Potato physiology. Li, P.H. Ed. Academic Press, Orlando. Pp. 1-34. Hudspeth, R.L. 1997: Cryptosporidiosis vaccine in edible plants. Crisp Data Base National Institutes of Health Agency, Project ID IR43AI41263-01. Jaswal, A.S. 1973: Effects of various processing methods on free and bound amino acid contents of potatoes. American Potato Journal 50: 86-95. Jaynes, J.M.; Derrick, K.S. Plants genetically engineered for nutritional quality. US Patent 5,811,654. Jaynes, J.M.; Langridge, P.; Anderson, K.; Bond, C.; Newman, C.W. 1985: Construction and expression of synthetic DNA fragments coding for polypeptides with elevated levels of essential amino acids. Applied Microbiology and Biotechnology 21: 200-205. Kaldy, M.S.; Markakis, P. 1972: Amino acid composition of selected potato varieties. Journal of Food Science 37: 375-377. Kikugawa, K.; Hakamada, T.; Hasunuma, M.; Kurechi, T. 1983: Reaction of p-hydroxycinnamic acid derivatives with nitrite and its relevance to nitrosamine formation. Journal of Agricultural and Food Chemistry 31: 780785. Page 34 Lang, S.L. 1992: Potato skins found to have chemical residues. Cornell Focus 1: 30. Langille, A.R.; Lan, Y. Gustine, D.L. 1998: Seeking improved nutritional properties for the potato: ethionine-resistant protoclones. American Journal of Potato Research 75: 201-205. Lazarov, K.; Werman, M.J. 1996: Hypocholesterolaemic effect of potato peel as a dietary fiber source. Medical Science Research 24: 581-582. Levander, O.A. 1990: Fruit and vegetable contributions to dietary mineral intake in human health and disease. HortScience 25: 1486-1488. Lewis, C.E. 1996: Anthocyanins and related compounds in potatoes (Solanum tuberosum L.). PhD Thesis, University of Canterbury, Christchurch, New Zealand. Lisiska, G.; Leszcyski, W. 1989: Potato science and technology. Elsevier Applied Science, New York. Lister, C.E. 1999: Natural antioxidants in fruit and vegetables. Leatherhead Food RA Food Industry Journal 2: 251-264. Lister, C.E.; Podivinsky, E. 1998: Health benefits of New Zealand vegetables. Prepared for Vegfed. FoodInfo Confidential Report 293. New Zealand Institute for Crop & Food Research Ltd. Lister, C.E.; Wilson, P.E.; Vile, G.F.; Sutton, K.H.; Simmonds, H.J. 2000: Pigments: not just a pretty face. Proceedings of the Nutrition Society of New Zealand 24: 34-39. Liu, M.A. 1999: Vaccines in the 21 st century. British Medical Journal 319: 1301. Ma, S.W.; Zhao, D.L.; Yin, Z.Q.; Mukherjee, R.; Singh, B.; Qin, H.Y.; Stiller, C.R.; Jevnikar, A.M. 1997: Transgenic plants expressing autoantigens fed to mice to induce oral immune tolerance. Nature Medicine 3: 793-796. McCay, C.M.; McCay, J.B.; Smith, O. 1987: The nutritive value of potatoes In: Potato Processing, Talburt, W.F.; Smith, O. (Ed.) Van Nostrand Reinhold, New York. Pp. 287-331. Markakis, P. 1975: The nutritive value of potato protein In: Protein nutritional quality of foods and feeds, Part 2; Friedman, M., Ed. Dekker, New York. Pp. 471-487. Mason, H.S.; Haq, T.A.; Clements, J.D.; Arntzen, C.J. 1998: Edible vaccine protects mice against Escherichia coli heat-labile enterotoxin (LT): potatoes expressing a synthetic LT-B gene. Vaccine 16: 1336-1343. Miller, D.L.; Castellanos, V.H.; Shide, D.J.; Peters, J.C.; Rolls, B.J. 1998: Effect of fat-free potato chips with and without nutrition labels on fat and energy intakes. American Journal of Clinical Nutrition 68: 282-290. Monro, J.A. 1999: Available carbohydrate and glycemic index combined in new data sets for managing glycemia and diabetes. Journal of the Science of Food and Agriculture 12: 71-82. Page 35 Morris, S.C.; Petermann, J.B. 1985: Genetic and environmental effects on levels of glycoalkaloids in cultivars of potato (Solanum tuberosum). Food Technology Australia 36: 118-124. Mullineaux, P.M.; Creissen, G.P. 1996: Opportunities for the genetic manipulation of antioxidants in plant foods. Biochemical Society Transactions 24: 829-835. Nestares, T.; Lopez-Jurado, M.; Sanz, A.; Lopez-Frias, M. 1993: Nutritional assessment of two vegetable protein concentrates in growing rats. Journal of Agricultural and Food Chemistry 41: 1282-1286. Oakes, J.V.; Shewmaker, C.K.; Stalker, D.M. 1991: Production of cyclodextrins, a novel carbohydrate, in the tubers of transgenic potato plants. Biotechnology 9: 982-986. Onyeneho, S.N.; Hettiarachchy, N.S. 1993: Antioxidant activity, fatty acids and phenolic acids compositions of potato peels. Journal of the Science of Food and Agriculture 62: 345-350. Oste, K. 1989: Impact damage, gangrene and dry rot in potato – important biochemical factors in screening for resistance and quality in breeding material. PhD Thesis. The Swedish University of Agricultural Sciences, Svalov. Pirie, N.W. 1973: The effects of processing conditions on the quality of leaf protein. In: Protein nutritional quality of foods, Vol. 2; Friedman, M. Ed. Dekker, New York. Pp. 341-354. Prince, D.M.; Welschenbach, M.A. 1998: Olestra: a new food additive. Journal of the American Dietetic Association 98: 565-9. Raben, A.; Andersen, K.; Karberg, M.A.; Holst, J.J.; Astrup, A. 1997: Acetylation of or beta-cyclodextrin addition to potato beneficial effect on glucose metabolism and appetite sensations. American Journal of Clinical Nutrition 66: 304-314. Rexen, B. 1976: Studies of protein of potatoes. Potato Research 19: 189-202. Rodriguez de Sotillo, D.; Hadley, M.; Wolf-Hall, C. 1998: Potato peel extract a nonmutagenic antioxidant with potential antimicrobial activity. Journal of Food Science 63: 907-910. Russell,D.G.; Parnell, W.R.; Wilson, N.C. et al. 1999: NZ Food: NZ People. Key results of the 1997 National Nutrition Survey. Ministry of Health, Wellington, New Zealand. Salmanian, A.-H.; Gushchin, A.; Medvedeva, T.; Noori-Daloii, M.R.; Domansky, N. 1996: Synthesis and expression of the gene for human epidermal growth factor in transgenic potato plants. Biotechnology Letters 18: 1095-1098. Salunkhe, D.K.; Wu, M.T.; Jadhav, S.J. 1972: Effects of light and temperature on the formation of solanine in potato slices. Journal of Food Science 37: 969-970. Page 36 Smith, G.; Walmsley, A.; Polkinghorne, I. 1997: Plant-derived immunocontraceptive vaccines. Reproduction, Fertilisation and Development 9: 85-89. Soh, N.L., Brand-Miller, J 1999: The glycaemic index of potatoes: the effect of variety, cooking method, and maturity. European Journal of Clinical Nutrition 53: 249-254. Somer, A.; West, K.P. 1966: Vitamin A deficiency: health, survival and vision. Oxford University Press, New York and Oxford. Spencer, C.M.; Cal, Y.; Martin, R.; Gaffney, S.H.; Goulding, P.N.; Magnolato, D.; Lilley, T.H.; Haslam, E. 1988: Polyphenol complexation – some thoughts and observations. Phytochemistry 27: 2397-2409. Stark, J.C.; Corsini, D.L.; Hurley, P.J.; Dwelle, R.B. 1985: Biochemical characteristics of potato clones differing in blackspot susceptibility. American Potato Journal 62: 657-666. Stich, H.F.; Rosin, M.P. 1984: Naturally occurring phenolics as antimutagenic and anticarcinogenic agents. In: Nutritional and Toxicological Aspects of Food Safety, Friedman, M., Ed. Plenum, New York. Pp. 1-29. Tacket, C.O.; Mason, H.S.; Losonsky, G.; Clements, J.D.; Levine, M.M.; Arntzen, C.J. 1998: Immunogenicity in humans of a recombinant bacterial antigen delivered in a transgenic potato. Nature Medicine 4:607-9. Thompson, L.U.; Yoon, J.H.; Jenkins, D.J.A.; Wolwer, J.; Jenkins, A.L. 1983: Relationship between polyphenol intake and blood glucose response of normal and diabetic individuals. American Journal of Clinical Nutrition 39: 745-751. Toma, R.B.; Augustin, J.; Orr, P.H.; True, R.H.; Hogan, J.M.; Shaw, R.L. 1978: Changes in the nutrient composition of potatoes during home preparation: I. Proximate composition. American Potato Journal 55: 639-645. True, R.H.; Hogan, J.M.; Augustin, J.; Johnson, S.R.; Teitzel, C.; Toma, R.B.; Orr, P. 1979: Changes in the nutrient composition of potatoes during home preparation: III. Minerals. American Potato Journal 56: 339-350. Tu, H.M.; Godfrey, L.W.; Sun, S.M. 1994: Expression of the Brazil nut methionine-rich protein in transgenic potato plants. In: The molecular and cell biology of the potato. 2nd ed. Belnap, W.R.; Vayda, M.E.; Park, W.D., Eds. CAB International, Wallingford, UK. Pp. 209-220. Vayda, M.E.; Belknap, W.R. 1992: The emergence of transgenic potatoes as commercial products and tools for basic science. Transgenic Research 1: 149-163. Vinson, J.A.; Hao, Y.; Su, X.; Zubik, L. 1998: Phenol antioxidant quantity and quality in foods: vegetables. Journal of Agricultural and Food Chemistry 46: 3630-3634. Vinson, J.A.; Jan, J.; Dabbagh, Y.A.; Serry, M.M.; Cai, S. 1995: Plant polyphenols exhibit lipoprotein-bound antioxidant activity using an in vitro Page 37 oxidation model for heart disease. Journal of Agricultural and Food Chemistry 43: 2687-2689. Weaver, M.L.; Timm, H.; Ng, H. 1983: Changes in nutritional composition of Russet Burbank potatoes by different processing methods. American Potato Journal 60: 735-744. Widholm, J.M. 1977: Selection and characterisation of amino acid resistant plant cell cultures. Crop Science 17: 597-600. Woolfe, J.A. 1986: The potato in the human diet. Cambridge University Press, Cambridge, UK. Pp. 58-82. Page 38 Appendix I - the chemical composition of New Zealand-grown potatoes, raw and cooked by a variety of method Other foods that may replace potatoes in a meal are also included for comparison. For details of analytical methods refer to The Concise New Zealand Food Composition Tables (Burlingame et al. 1997). Page 39 Water Energy Energy Protein (Nitrogen x 6.25) Total fat Carbohydrate, available Dietary fibre (Englyst) Ash Table 8: Proximate composition. g kcal kJ g g g g g Combined cultivars, flesh and skin, raw 78.54 72 299 2.36 0.11 15.5 1.3 1.15 Combined cultivars, flesh, raw 78.17 70 291 2.06 0.06 15.4 1.4 0.97 Fries, fried in beef dripping, salt added 56.44 186 773 3.77 6.26 28.7 1.4 1.69 Fries, fried in butter, salt added 63.33 159 660 2.92 4.71 26.3 1.4 1.65 Fries, fried in peanut oil, salt added 58.18 174 723 3.46 5.34 28.1 1.4 1.35 Fries, frozen, uncooked 73.1 107 445 2.2 3 17.9 1.3 0.6 Fries, independent shops, ready to eat 52.9 241 1002 3.8 14 24.9 1.6 2 Ilam Hardy, flesh, raw 79.07 68 279 1.97 0.05 14.8 1.63 0.93 In skin, microwaved 77.9 74 305 1.94 0.07 16.34 2.7 1.05 Instant powder, prepared with water 79.4 69 283 2 0.2 14.7 1.2 1.1 81 72 297 2.36 0.06 15.5 1.3 1.15 Red King, flesh, raw 79.6 70 291 2.06 0.06 15.4 1.2 0.97 Rua, flesh, baked, salt added 75.36 88 366 2.58 0.15 19.2 1.99 1.28 Rua, flesh, boiled, drained 77.06 83 342 2.1 0.17 18.2 1.73 0.88 Rua, flesh, boiled, mashed with milk, butter and salt added 79.33 95 394 1.84 3.29 14.5 1.7 Rua, flesh, boiled, salt added 77.06 83 342 2.1 0.17 18.2 1.73 0.88 Rua, flesh, microwaved, salt added 72.39 95 394 2.48 0.13 21.1 2.3 Rua, flesh, raw 79.91 70 291 1.78 0.15 15.5 1.56 0.79 Rua, flesh, roasted in beef dripping, salt added 71.82 105 433 2.46 0.64 22.3 1.42 1.43 POTATOES Red King, flesh and skin, raw Page 40 0.73 1.34 Dietary fibre (Englyst) Carbohydrate, available kcal 73.2 kJ 306 g g 2.38 0.111 g 15.7 g g 1.31 1.16 Bread, white, sliced 39 216 904 7.3 0.9 43.4 2.8 Bread, wholemeal 45 198 829 8.1 1.4 37.1 5.7 Macaroni, boiled 78 86 359 3 0.5 16.8 0.9 Rice, brown, boiled 66 141 589 2.6 1.1 29.2 1.8 Rice, white, polished, boiled 70 123 513 2.2 0.3 26.9 1.3 Spaghetti, boiled 74 100 420 3.6 0.3 20.1 1.2 OTHER Page 41 Ash Energy Total fat Energy Protein (Nitrogen x 6.25) Water g 78.3 Whole, with skin, fried Sulphur Chloride Potassium mg 35 mg 55 mg 444 2.54 22.9 46.1 32 55 145 33 69 76 Fries, fried in butter, salt added 158 27 57 Fries, fried in peanut oil, salt added 179 31 Fries, frozen, uncooked 25 21 Fries, independent shops, ready to eat 250 27 Ilam Hardy, flesh, raw In skin, microwaved Instant powder, prepared with water Selenium Fries, fried in beef dripping, salt added Zinc Combined cultivars, flesh, raw Copper mg 17 Iron mg 4 Manganese Combined cultivars, flesh and skin, raw Calcium Magnesium mg 33 Sodium Phosphorus Table 9: Nutrient elements. mg 3 µg 115 mg 0.6 mg 0.093 mg 0.29 µg 1.05 484 4.39 106 0.57 0.186 0.2 0.27 223 881 7 235 1.3 0.25 0.6 0.35 60 244 776 8 176 0.7 0.19 0.4 1.2 69 70 276 770 6 238 1 0.27 0.6 0.35 61 32 45 420 8 169 0.7 0.11 0.3 0.27 70 40 65 650 11 146 1.3 0.12 0.6 0.35 2.43 21.9 44.2 30.7 52.8 464 4.21 161 0.55 0.209 0.39 0.27 1.6 23 48 220 0.83 0.12 0.27 T POTATOES 42 - 510 6.7 260 15 48 6 380 340 20 79 0.5 0.08 0.2 0.27 Red King, flesh and skin, raw 3 17 5 32 55 484 4 116 0.57 0.086 0.23 0.25 Red King, flesh, raw 3 17 4.6 32 55 484 4 68.2 0.57 0.131 0.22 0.249 104 18 41 40 160 543 4 116 0.3 0.14 0.3 0.98 4 15 32 35 26 332 4 102 0.5 0.15 0.2 0.47 Rua, flesh, boiled, mashed with milk, butter and salt added 130 12 39 30 265 282 21 70 0.3 0.13 0.3 0.4 Rua, flesh, boiled, salt added 127 15 32 35 318 332 4 102 0.5 0.15 0.2 0.47 Rua, flesh, microwaved, salt added 119 22 47 46 183 610 4 134 0.5 0.18 0.4 0.98 Rua, flesh, baked, salt added Rua, flesh, boiled, drained Rua, flesh, raw 4 17 33 35 55 444 3 120 0.57 0.14 0.3 0.27 Rua, flesh, roasted in beef dripping, salt added 124 22 44 47 191 589 6 138 0.6 0.17 0.3 0.35 Whole, with skin, fried 4.04 17.2 33.3 - - Page 42 448 3.03 120 0.606 0.0939 0.293 1.1 Sodium Magnesium Phosphorus Sulphur Chloride Potassium Calcium Manganese Iron Copper Zinc Selenium mg mg mg mg mg mg mg µg mg mg mg µg OTHER Bread, white, sliced 666 162 40 1 0.8 4 Bread, wholemeal 641 227 33 1.7 1.3 3.2 Macaroni, boiled 1 25 5 0.4 0.5 0.3 Rice, brown, boiled 1 99 4 0.5 0.7 2.3 Rice, white, polished, boiled 2 38 1 0.2 0.1 2.6 Spaghetti, boiled T 24 5 0.4 0.5 T Page 43 Retinol Beta-carotene equivalents Total vitamin A equivalents Thiamin Riboflavin Niacin Vitamin B6 Pantothenate Biotin Folate, total Vitamin B12 Vitamin C Vitamin D Alpha-tocopherol Vitamin E Table 10: Vitamin composition. µg µg µg mg mg mg mg mg µg µg µg mg µg mg mg 0 6 1 POTATOES Combined cultivars, flesh and skin, raw 0.09 0.04 1.474 0.07 0.38 0.1 15 0 19 0 0.07 0.08 0.086 0.003 0.887 0.025 0.38 0.1 14 0 12.4 0 0.073 0.083 0.2 0.4 10 T 11 0 0.34 0.09 13 T 10 0.03 - Combined cultivars, flesh, raw 0 6 1 Fries, fried in beef dripping, salt added T 34 6 0.13 0.1 0.9 0.15 Fries, fried in butter, salt added T 71 12 0.13 0.04 0.4 0.13 Fries, fried in peanut oil, salt added 0 23 4 0.14 0.05 0.5 0.14 0.487 0.4 11 0 4 0 Fries, frozen, uncooked 0 T T 0.08 0.01 1.6 0.28 0.42 0.1 12 0 6 0 0.073 0.083 Fries, independent shops, ready to eat 0 0 0 0.1 0.03 1.1 0.27 0.498 0.4 9 0 - Ilam Hardy, flesh, raw 0 0 0 In skin, microwaved 0 T T Instant powder, prepared with water 0 54 Red King, flesh and skin, raw 0 6 Red King, flesh, raw 0 Rua, flesh, baked, salt added Rua, flesh, boiled, drained 0.082 0.003 16 0 0.85 0.024 0.39 0.1 14 0 11.9 0 1.04 0.1 0.46 0.46 44 0 0 - 1.2 0.1 0.2 0.1 0.1 0.36 0.07 0.08 0.04 T 9 0.01 0.03 0.18 0.2 0.3 5 0 3 1 0.086 0.03 0.89 0.025 0.38 0.1 15 0 12 0 0.073 0.083 6 1 0.086 0.03 0.89 0.025 0.39 0.1 14 0 12 0 0.073 0.083 0 7 1 0.09 0.04 0.5 0.09 0.34 0.09 13 0 10 0 0.073 0.083 0 7 1 0.07 0.04 0.3 0.07 0.34 0.07 13 0 9 0 0.073 0.083 Rua, flesh, boiled, mashed with milk, butter and salt added 5 39 11.5 0.05 0.05 0.5 0.07 0.33 0.25 12 0.01 Rua, flesh, boiled, salt added 0 7 1 0.07 0.04 0.3 0.07 0.34 0.07 13 0 9 0 0.073 0.083 Rua, flesh, microwaved, salt added 0 13 2 0.09 0.05 0.3 0.1 0.34 0.09 13 0 10 0 0.073 0.083 Rua, flesh, raw 0 6 1 0.07 0.04 0.4 0.07 0.38 14 0 12 0 0.073 0.083 Page 44 0.1 1.5 0.1 0 8 0.01 0.11 0.02 0.021 0.09 0.1 0 0.33 0.09 3.3 Bread, wholemeal 0 0 0.66 0.16 Macaroni, boiled 0 0 0.06 0.02 Rice, brown, boiled 0 0 0.14 Rice, white, polished, boiled 0 0 Spaghetti, boiled 0 0 mg µg µg µg mg µg mg mg 0.34 0.09 13 0 10 0 0.075 0.085 - 11 0 19 0.02 28 0 0 3.9 0.04 36 0 0 0.7 0.01 3 0 0 0.02 1.9 0.19 10 0 0 0.01 0.01 0.8 0.05 3 0 0 0.02 0.01 0.8 0.02 4 0 0 OTHER Page 45 Vitamin E 0 Alpha-tocopherol Bread, white, sliced 1.4 0.067 - Vitamin D 0.073 0.038 Vitamin C 1 Vitamin B12 6.1 mg 0.11 Folate, total 0 mg 0.8 Biotin Whole, with skin, fried mg 0.04 Pantothenate mg 0.11 Vitamin B6 µg 2 Niacin Total vitamin A equivalents µg 10 Riboflavin Beta-carotene equivalents µg T Thiamin Retinol Rua, flesh, roasted in beef dripping, salt added 0 - 0.081 Isoleucine Leucine Lysine Methionine Cysteine Phenylalanine Tyrosine Threonine Tryptophan Valine Arginine Histidine Alanine Aspartic acid Glutamic acid Glycine Proline Serine Table 11: Amino acid composition (g per 100 g FW). Combined cultivars, flesh and skin, raw - - - - - - - - - - - - - - - - - - Combined cultivars, flesh, raw - - - - - - - - - - - - - - - - - - 0.158 0.227 0.208 0.06 0.048 0.168 0.119 0.148 0.054 0.198 0.188 0.072 0.139 0.693 0.484 0.129 0.148 0.158 - - - - - - - - - - - - - - - - - - Fries, fried in peanut oil, salt added 0.15 0.207 0.184 0.055 0.044 0.15 0.105 0.138 0.05 0.173 0.173 0.067 0.127 0.634 0.438 0.115 0.138 0.15 Fries, frozen, uncooked 0.091 0.13 0.12 0.035 0.028 0.095 0.067 0.084 0.032 0.11 0.11 0.042 0.081 0.4 0.28 0.074 0.084 0.091 Fries, independent shops, ready to eat - - - - - - - - - - - - - - - - - - Ilam Hardy, flesh, raw - - - - - - - - - - - - - - - - - - In skin, microwaved - - - - - - - - - - - - - - - - - - 0.083 0.12 0.11 0.032 0.026 0.086 0.061 0.077 0.029 0.1 0.099 0.038 0.074 0.37 0.26 0.067 0.077 0.083 Red King, flesh and skin, raw - - - - - - - - - - - - - - - - - - Red King, flesh, raw - - - - - - - - - - - - - - - - - - Rua, flesh, baked, salt added 0.11 0.161 0.141 0.041 0.043 0.111 0.078 0.099 0.037 0.131 0.131 0.05 0.095 0.473 0.332 0.087 0.099 0.111 Rua, flesh, boiled, drained 0.088 0.127 0.114 0.034 0.026 0.091 0.064 0.08 0.031 0.108 0.104 0.041 0.077 0.395 0.263 0.07 0.08 0.088 Rua, flesh, boiled, mashed with milk, butter and salt added 0.076 0.111 0.1 0.029 0.023 0.08 0.056 0.071 0.027 0.094 0.09 0.036 0.067 0.343 0.233 0.061 0.071 0.076 Rua, flesh, boiled, salt added 0.087 0.128 0.114 0.034 0.027 0.091 0.065 0.081 0.031 0.108 0.105 0.04 0.078 0.39 0.269 0.071 0.081 0.087 - - - - - - - - - - - - - - - - - - Rua, flesh, raw 0.074 0.109 0.1 0.028 0.022 0.077 0.054 0.069 0.026 0.092 0.092 0.034 0.065 0.326 0.226 0.059 0.069 0.074 Rua, flesh, roasted in beef dripping, salt added 0.105 0.149 0.131 0.039 0.032 0.105 0.075 0.096 0.036 0.122 0.122 0.047 0.087 0.455 0.315 0.083 0.096 0.105 - - - - - - - - - - - - - - - - - - POTATOES Fries, fried in beef dripping, salt added Fries, fried in butter, salt added Instant powder, prepared with water Rua, flesh, microwaved, salt added Whole, with skin, fried Page 46 Edible portion g W eight Common Measure W eight Common Measure Table 12: Common serving sizes and edible portions. g % POTATOES Combined cultivars, flesh and skin, raw 1 potato 150 Combined cultivars, flesh, raw 1 potato 117 96 Fries, fried in beef dripping, salt added 1 cup 60 Fries, fried in butter, salt added 1 cup Fries, fried in peanut oil, salt added 1 cup Fries, frozen, uncooked 10 chips 65 100 Fries, independent shops, ready to eat 1 cup 100 100 Ilam Hardy, flesh, raw 1 potato 117 85 Instant powder, prepared with water 1 cup 241 100 Red King, flesh and skin, raw 1 potato 150 99 Red King, flesh, raw 1 potato 117 88 Rua, flesh, baked, salt added 1 potato 90 1 cup 128 100 Rua, flesh, boiled, drained 1 potato 114 1 cup 164 100 Rua, flesh, boiled, mashed with milk, butter and salt added 1 cup 209 Rua, flesh, boiled, salt added 1 potato 114 1 cup 164 Rua, flesh, microwaved, salt added 1 potato 90 1 cup 128 Rua, flesh, raw 1 potato 117 Rua, flesh, roasted in beef dripping, salt added 1 potato 95 1 cup 130 Whole, with skin, fried 1 potato 97 1 cup 134 Bread, white, sliced 1 medium slice 26 1 thick slice Bread, wholemeal 1 medium slice 28 Macaroni, boiled 1 cup 149 Rice, brown, boiled 1 cup 206 Rice, white, polished, boiled 1 cup 216 Spaghetti, boiled 1 cup 148 1 cup, diced 158 84 10 chips 45 100 60 10 chips 45 100 60 10 chips 45 100 In skin, microwaved 100 100 84 100 OTHER Page 47 36 100