View of Sustainability - a challenge to animal production and breeding

Review

Sustainability

^-

^a challenge ^to anima production ^and breeding

Heidi Torp-Donner, Jarmo Juga

DepartmentofAnimal Science, FIN-00014 Universityof^Helsinki,Finland. Current address: Finnish Animal Breeding Association,^POBox40, 01301 Vantaa, Finland,e-mail:heidi.torp@mloy.fi

Prospects of sustainable animal breeding are briefly reviewed from the animal breeding point of view. The aspects ofsustainabilityincluded are:ecological sustainability suchasenvironmental^sus- tainabilityandmaintenanceofbiodiversity aswellasethical and economicalsustainability.Environ-

mentaldegradationcan^bereducedbyintensiveproductionatleastonintermediateproduction levels.

Biodiversity of livestock breedscan be maintained withgloballydiverse breeding goals containing several traits and with national conservation schemes forrarelocal breeds. Ethicalsustainabilitycan be taken into accountby improvinghealth andlongevity traits. Production must also be economically profitable,otherwise it is not sustainable.Inoptimisingall these aspects, the animal breeders have to laydown criteria for conservation programmes and re-evaluate breeding goals so thatsustainability is taken into account.

Key ^words', biodiversity, ecological, economical,environmental sustainability, ethical, sustainable

livestockproduction

ntroduction

Sustainable livestock production means a pro- duction that is ecologically, taking intoaccount the environment and biodiversity, ethically and economically sustainable. Noconcise,universal- ly acceptable definition of sustainable agricul- ture has yet emerged, but most agriculturists agree that theconceptof sustainable agriculture is ofparamount importance^tothe sustainability ofour biosphere and its ever-increasing human population (Heitschmidt etal. 1996).

Sustainable agricultural _systems should maintain _or enhance environmental quality,use non-renewable_resources wisely, _promote the maintenance of renewableresources including animal and plant biodiversity andpromote eco- nomic viability. AccordingtoVavra(1996) sus- tainablesystems arethose that exist in theover- lap of what the^current generation wantsfor it- self and future generations and what is biologi- cally and physically achievable in the longrun.

Inlivestock production, sustainability could meanbeing abletoharvest thesamequantity of meat, milk orfibre from a given land base in-

©Agricultural and Food ScienceinFinland ManuscriptreceivedMay 1997

Torp-Donner, H. ^& Juga, J. Sustainability^- animal production and breeding definitely (Vavra 1996 and Heitschmidt _et al.

1996).Because of the large heterogeneity of live- stock production systems, asystem-specific anal- ysis is needed to evaluate all these criteria for sustainability.

Research and extension services needtopro- vide leadership in developing alternativesystems that provide the broad view of sustainability be- forewe arepainfully forced into it by restrictive legislation (Vavra 1996).An environmental pro- gramme ofagriculture resulting from the regu- lations of the European Union has already been prepared by the Ministry ofAgriculture and For- estryin Finland (The Ministry ofAgriculture and Forestry 1995). The requirements listed in the programme will lead ^to substantial investment costs for farmers and the effects of thesecosts on livestock production havenotbeen studied.

The goal of this review is todiscuss the fol- lowing criteria thatareusedtodescribe sustain- able livestock production: 1) the environment:

air, surface and groundwater quality and degra- dation,2)the biodiversity oflivestock,its meas- ures and the opportunities and dangers of bio- technology, 3) the ethical aspects of livestock production that consist of human attitudes and the welfare and health ofanimals, and 4) the economical aspects of the three previous crite- ria. Animal breeding methods that could enhance

sustainable livestock production are also dis- cussed. Future research for defining and evalu- ating sustainability will also be suggested.

Environment

Air quality

Emissions from agriculture into the air include methane(CH₄^),carbon dioxide(C0₂⁾and nitrous oxide(N₂0),which speed up climate change,as well_asammonia (NH ^),which_causeseutrophi- cation of the soil and water. Agriculture, espe- cially livestock production, is the main source of methane emissions in Finland ^(55% of all

methane emissions). Emissions from othersources are not so great (Yläranta 1991). Emission of methane, carbon dioxide and nitrous oxidecan be reducedtosomeextent by new methods and technologies for manure and urine storage and handling.

The major cause of methane emission ^is, however,ruminant digestion. That emissioncan be diminished by reducing the number of ani- mals and by optimising the feeding of animals.

On the other hand, if food production needs_to be maintained_atthe present level, onepossible waytodiminish methane emission istoincrease the production efficiency per animal. Calcula- tions show that methane emissionsarereduced from30 g/kg of milk produced^to 15 g/kg when annual milk production is increased from4 000 kg ofmilk per animalto8 000 kg (Flachovsky 1994).

Surface and ground water quality

Phosphorus and other nutrient loads in surface wateris a major problem in Finland because of rainyautumnsand melting snow in spring. Nu- trient loads in surface water causeeutrophica- tion. Over 80% of arable land in Finland is in grass production, which prevents the nutrient load from reaching the surface^water.Livestock production is thus superior to cropproduction from the nutrient load point of view(TheMinis- try of Agriculture and Forestry 1995). Nutrient load in surfacewateraswellasin ground water can be diminishedby improved cultivation meth-

ods and by increasing grass production.

Preliminary calculations showareduction in N-emissions from 337 g/kg ofmeatproduced^to 231 g/kg, when the daily growth ofabeef ani- mal is increased from 800 g/dayto 1600 g/day (Korhonen 1996). It is also estimated that the improvement of the production performance of animals realised in the last decade by improved breeding programmes has contributed^toreduce the polluting power by, approximately 25% in swine and 30% in poultry (Matassino etal.

1991).These resultssuggest that increased effi- ciency is environmentally moresustainable than

Table I.^{Total N-andP-}emissions (kg) per 1million kg product (meat^ormilk) and returns (FIM per kg N-orP-^emission)indifferentproduction levelsinmeatandindairy production.

Meatproduction (g/day) Milkproduction (kg/year)

8(K) 1200 1600 4000 6000 8000

Return(FIM/year)" 3 998 5 998 7 997 7 840 11760 15 680

N(kg/1 mill.kg prod.) ^2>326 300 243 100 224 000 "20000 16 000 15 000

Return(FIM/N-kg) 43 57 62 98 123 131

P(kg/1 mill.kg prod.) ^2»66 000 55 600 57 300 "3 000 2 000 1500

Return(FIM/P-kg) 210 250 242 653 980 1307

•>Evaluated from the basis of the mean producer prices inyear 1995 paid inFinland (Maaseudun

Tulevaisuus 23.3.1996).

2>Evaluated from the basis of calculations about feed costs andN-and P-emissionsinmeatproduction (Korhonen 1996).

3)Evaluated from the basis of calculations madeby Flachowsky 1994.

extensive goals. This is certainly thecasefor low and intermediate production levels,butmorein- formation about environmental efficiency is still needed for very intense production ^systems. A very intense beefproduction system, which has a heavy reliance on fossil fuels and ^extrapro- tein foodstomaintain aproductivecow herd in regions where nutrient shortfallsare common, carries with it some ecological and economic risks (Heitschmidtetal. 1996).These risks arise from the assumption of the availability of cheap sources of exogenous energy and the potential disruption of critical life-supporting ecological systems due to continued generation of de- gradants (Heitschmidtetal. 1996).

InTable 1 the incomes (FIM) are evaluated perkg of N- and P-emissions from manureand urine for dairy cows and for meat production.

Dairy cows bring in twiceas much money per kg of N emittedasdomeat animals and the dif- ference in P-emission isevenbigger. Thus milk production could bettercoverthecoststhat arise when accounting for environmentalaspects.The calculations also show that the financial incomes are proportionally higher when the production increases fromalow production level^toan in- termediate level than fromanintermediate level toahigh production level. The P-emissions pro- duced by meatproduction turnedout tobeeven

higher in high production levels than in inter- mediate production levels.

Biologically sustainable husbandry systems for cattle and sheep canbe achieved by sustain- able nutrientmanagement.This implies finetun- ing of nutrient(N, P andK) input and output ^to achieve and maintain agronomically desired and ecologically acceptablereserves of nutrients in the soil(Hermans and Vereijken ^1994).The an- nual in- and output balance sheets of various nutrientsat afarm orregional level should be maintained and the available soil reserves of nutrients should also be evaluated. Nutrient flows are asfollows: nutrient inputtoanagri-environ- mental system comesfrom fertilisers,feeds and biological nutrient fixation and nutrientoutput contains products and ^manure, if transported from thefarm, losses as nutrientvolatilisation, run-off and leaching ^(Hermans and Vereijken

1994).

Selection of livestock for efficiency of feed utilisation resulted in decreased losses in the poultry meatproduction cycle through manure, slaughter offal and mortality. Feed costs were also reduced and the slaughter yield wasequiv- alentor improved when compared with selec- tion for growth ^rate (Leenstra and Ehlhardt

1994). These direct and correlated effects of_se- lection of livestock_onthe efficiency of feed uti-

Torp-Donner, H. ^& Juga, J. Sustainability^- animal production and breeding lisation indicate good means for establishing

environmentally acceptable poultry meatproduc- tion. Two _aspects did_not favour the selection for efficiency of feed utilisation: the increased rearing period and the increasedcostsof these-

lection itself (Leenstra and Ehlhardt ^1994).How- ever,if environmental degradation is includedas

anextracost,it will make themorecostly selec- tion of livestock profitable.

Another important measure of production efficiency is the efficiency of converting vege- table energy and protein intomeatormilk (Oltjen and Beckett 1996). Dairy production efficiency ranges from 96% to 276% on the basis of pro- duction of humanly consumable protein. The protein resulting from ruminant livestock pro- duction is of higher quality withgreaterbiolog- ical value than the protein in the substrate feeds (Oltjen and Beckett 1996)Thus theargumentthat ruminant livestock belong in sustainable live- stock production is convincing.

Biodiversity

Biodiversity ofall_ecosystems stabilises the func- tions of thesesystemsand their interactions with surroundingecosystems. Thus, agri-ecosystems influence their surroundings and viceversa.Sus- tainable management of diversity in livestock production aims for the sustainableuse of ge- netic_resources and the maintenance of genetic variability for present and future needs. The Finnish agri-environmentalprogramme pays pre-

miums for preserving traditional biotopes and rare local breeds (The Ministry of Agriculture and Forestry 1996),which has sometimes been justified by the future needs of animal breeding, but has more ^todo with conserving the Finnish cultural heritage. The agri-ecosystem accounts for_a small proportion of total biological diver- sity, but impact on human survival is of great importance. Thismeans that agricultural biodi- versity enables continuous food production for man in differentenvironments, but has little

meaning if the evolution of all ecological biodi- versity is considered (Haila 1995). Specific ag- ricultural environments around the world impose biological limitations for the production ^animals, and thus breeding goals for specific environ- ments are needed, which on the otherhand, en- hance global biodiversity (Beilharzetal. 1993).

New mutations

Variability between and within different species and breeds is caused by evolutionary forces like mutation, naturalselection, isolation, migration and random drift. In proteins, neutral mutations are morefrequent than theoneswhichare strong- ly selected against. Mutations providenewvari- ation,and thus newresponsein selectioncanbe achieved. The effects of mutationsare usually smallor neutral, but the balance between muta- tions and selection the responsetoselection has been achieved in long-term selection pro- grammes,asthe number of mutative loci isgreat in quantitative traits. Mutations with large ef- fects can cause newachievements_orchanges in breeding programmes (Hill 1989).Well-known examples of major genes in animal breedingare:

the halothane gene in swine, the dwarf gene in poultry, the double muscling gene in cattle and the boorola gene in sheep (Mackay 1989).Many of these major genes have negative effects on viability and fertility. Mutations with negative effects on fitness are quickly eliminated. Thus variability caused by mutations is greater, the more neutral mutations are involved (Crow 1986). Nearly all mutationsarelost inafew gen- erations through random genetic driftevenif they were favourable, because their frequencies are low (mutation frequency ^/locus ^/generation ⁼ IOMO'^8).

Maintenance of variation in breeding populations

Selection decreases the genetic variance in breeding populations. The loss of genetic varia-

tion dependsonthe intensity of the selection of theparents of thenextgeneration, the heritabil-

ity, the population size and the mating design (Falconer 1989). Even the use of modern pre- diction methods like the animal model BLUP accelerates the reduction in genetic variation due to within-family selection (Meuwissen and Woolliams 1994^b). Use of marker-assisted se- lection might also result in a lower long-term

selection response due^toreduced genetic varia- bility (Gibson 1994).

The population size is very important in maintaining biodiversity. When population size decreases, the inbreedingrate increases asdoes homozygosity. The increased inbreeding_rate leadstoareduction in heterozygosity andtoin- creased homozygosity in the population, which

means decreased genetic variability. Heterozy- gosityas ameasurementof variability describes towhatextentvariability remains in the popula- tion (Crow 1986). Heterozygosity in one locus can be calculatedas:

Het⁼ 1

-X"'

sl ^pr,^>2,

where p is the frequency ofanallele in alo- cusand m is the number of different alleles. The meanofheterozygosity in different loci indicates the genetic variability withinabreed. Heterozy- gosity decreases in each generation by M 1N_e⁽N_e

=effective population size) (Crow 1986).In large populations mutations balance geneticfixation, but in small populations a random increase in homozygosity is _more probable. Hence breed- ers should_ensure that the effective population size remains large in ordertoavoid losing varia- tion by random drift (Dempfle 1990).

When the inbreedingrate increases,function- al traits suchasreproduction and fitness decrease 3-5% per 10% increase in the inbreeding coef- ficient (Cunningham 1995). However, the de- creasein functional traitscan also be duetoneg- ative genetic correlations with the traits in breed- ing goals. On the otherhand,fitness in apopu- lationcan increase by natural selection that fa- vours moreviable individuals. In the absence of correlated responses duetoartificial selection, the critical population size,atwhich the increase

in fitness due to natural selection and the de- crease due toinbreeding depression _are in bal- anceis approximately: D I 2a_wa^, where D ⁼the inbreeding depression with complete inbreeding and o ⁼ the additive genetic variance of fit- ness(Meuwissen and Woolliams 1994^a).

The correlations between quantitative pro- duction traits and the fitness traits affect there- sponsetoselection in the long^term. If fitness is high atan optimum level of production traits, the selection for fitness will maintain variability in production traits in the longterm (Hill 1989).

If fitness declines due^to a correlated negative responseto artificial selection, then a large in- crease in the critical population size is needed.

However if the negative response is larger than

theresponse tonaturalselection, areduction in fitnesscannotbe prevented. Effective population sizes that_{prevent a}decline in fitnessareusually greater than those which maximise the genetic gain of production efficiency,_sothe former is _a more stringent restriction _on effective popula- tion size (Meuwissen and Woolliams 1994^a).

Another requirement for population size is set by the needtoreduce uncertainties in predicting the changes from the selection applied ^(Meuwis- sen and Woolliams

1994

^b).

The between-breed variation in quantitative traits is often overlapping, whichmeansthat the breeds arealike. Therefore it is difficult ^to use quantitative traits todetermineexact differenc- esbetween the breeding populations and hence

todefine criteria for conservation purposes. This means that small populations with overlapping variation could be jointly used as a breeding population with lower risk of inbreeding and better chances to compete with other breeding populations.

Since future needs cannotbe predicted, the easiest way to maintain variability is to main- tain pure breeds either with in situ or ex situ methods. The best waytodo this istomaintain several competitive breeds in production, be- cause exsitu maintenance and the introgression for future needs ismoreexpensive and timecon-

suming than the maintenance in situ (Smith 1984).

Torp-Donner, H. ^&Juga,J. Sustainability^-animal production and breeding

Implications ^to breeding programmes

A practical question in any conservation pro- gramme is the level of diversitytobe maintained.

How much of the diversity hastobe maintained within a country, or does global diversity be- tween the different countries satisfy the neces- sary level of diversity? In addition ^tothepoten- tial loss ofbreeds,therecanbeapotentially high loss of genes withinabreed. In^cattle,for_exam- ple, this has been accelerated bynew techniques suchasartificialinsemination,including frozen semen,multiple ovulation and embryo transfer, which makes it possible tousethesameanimals in many countries, and new genetic evaluation methods as the animal model BLUP (Hodges

1991).

The aim in conserving local andrare breeds is complex. The importance of conservation of such breeds for future animal breeding is diffi- culttoforecast and involves highcostscompared tothe possible gains. The original breeds should be maintained because of their possible special traits in production or, asfor example in devel- opingcountries, the adaptation to local climate orbetter resistance to local diseases. More im- portant to many countries is, however,the cul- tural heritage. Thismeans that globalconserva- tion programmes will have problems in defining which conservation programmes should be fi- nanced.Hencethe funding of these programmes would be more logical on a national basis and global funds should be reserved for programmes in developing countries.

Rapid changesare taking place for example in the black and white dairy cattle populations due to the impact of the North American Hol- stein. In theFinnish Friesian population, the pro- portion of Holstein genes has increased during thepast tenyears. In Friesiancowsborn in 1980, the proportion of Holstein genes^was only 2%, but incowsborn in 1990 the proportionwas31%.

In Finnish Friesian bulls the proportion of Hol- stein geneswas26.5% and 56.8%, respectively.

In other Finnish dairy breedssomechanges have also occurred in breed contribution, but the changes have been much smaller than in Finn-

ish Friesian cattle (Lidauer, personal communi- cation).

The management of genetic variability in Finnish dairy breeds is monitored by Al-associ- ations freezing sperm in long-termstorage.How- ever,the conservation of genetic material isnot explicitly included in the breeding programme.

The national breeding goals maintaingenet- ic variability withina breed if the goals differ between countries. For example theuse of Hol- stein breed has spread ^tomanycountries, but if the breeding goals are not the same, the differ- entlines of Holstein will increase the diversity between the countries. Diverse national breed- ing goals which include many traits should main- tain genetic diversity both globally and locally, although this hasnot been quantified in anyre- searchreport todate.

Goddard(1992) has shown that if thegenet- ic correlation between the breeding goals in two countries is 0.60 at most, countries tendto se-

lect different bull sires. A pilot studyon Finnish Ayrshire bulls gaveacorrelation of0.84 between the total merit indices in Sweden and Finland, 0.66 between Norway and Finland and 0.84 be-

tweenNorway and Sweden. The Nordic coun- tries share apreference for breeding for total economic merit which includes yield traits and functional traits. The correlations suggest that some common bull sires could be used in Nor- dic breeding, but that the breeding goals arefar from a consensus.

The totalrate of genetic gain increasescon- siderably by increasing population size. Com- bining all Nordic red dairy cattle populations would increase the totalrecorded population size to757 800, the total performance testing capac- ity to 1 110 and the total number of young bulls progeny testedto 450 (Lindhé 1995)and hence improve themanagementof geneticresourcesin red breeds.

in pigs, the genetic process for reducing fat and increasing thepercentageof lean meat has been rapid in many countries and has been ac- companied by problems in _meat quality and

stresssusceptibility, especially insomeLandrace populations in which the halothane gene has ap-

proached the frequency of 0.9 (Hodges 1991).

In the Finnish swine population, the eradication of halothane gene _was very ^effective, and the gene was culled rapidly from both Finnish pig breeds, Landrace and Yorkshire (Puonti and Schulman 1988).The animal model BLUP -eval- uations have been calculated for fertility traits (Nylander and Mäntysaari 1991)and for produc- tion traits (Haltia etal. 1993),butmoreresearch is needed toachieve sustainable breeding goals.

Similar selection goals for both Finnish Landrace and Finnish Yorkshire have already increased the similarity of these breeds.

Ethical aspects

Human attitudes

Ethicalaspects of livestock production methods should be evaluated together with technical and economical _aspects before including them in animal production. The ethical evaluation done byconsumers is, however,based_onattitudes and

not onresearch facts. In acontingent evaluation research study (Siikamäki, personal communi- cation) _on the willingness ofconsumers topay more for the decrease inuseof pesticides in ag- riculture, it was concluded thatconsumers are willing to pay a somewhat higher price for a producttolower theuseof pesticides inFinland, but they donotask foratotal banonpesticides.

In another study (Sihvonen 1993)itwasconclud- ed thatconsumers arereadytopay somethingto improve animal production fromanethical point of view. Organic products, which also have high- erprices, have been selling quite well, so some consumers are acting on the basis of their atti- tudes.However,in the longrundifficulties might arise in marketing organic products with higher prices, if their supply increases.

It is also ^not clear thatconsumers will ac- ceptgenetic manipulation in food products. The ethical aspectsofnewbreeding techniques like marker-assisted selection and gene transfer

should also be included in future breeding goals.

That is why optimised production, knownas“In- tegrated Production” (El Titi _et al. 1993),will probably be the futuretrend, when considering allaspects affecting sustainable production.

The welfare and health of production animals

Breeding methods for sustainable and healthy animals which have genetic resistancetoproduc- tion-related diseases should be taken intocon- sideration more in future breeding goals, when production levels increase. Yield traitsarein fact genetically negatively correlatedtofertility traits (Pösö and Mäntysaari

1996

b) and udder health (Pösö and Mäntysaari 1996^a).New techniques of marker-assisted selection might offer possi- bilities for selection based on traits that affect production sustainability or production-linked disease resistance especially in traits that have low heritability.

In research concerning breeding and animal welfare, the economic influence of health and diseases should be evaluated inmore detail.The economic, aswellasthe ethical impact ofapro- duction related-disease is always negative, so breeding goals for animal health also ^warrant moreresearch in the future.

Economic aspects

Costs and premiums ^{in Finland}

Modelling sustainability economically, ecolog- ically and ethically forces us to evaluate^costs and benefits froma new perspective compared to previous production methods. The environ-

mental and ethical aspects are very difficult^to evaluate in monetary terms, but they should be taken into account in the modelling of produc- tion which optimises environmental andeconom-

icalaspects in every production scheme(ElTiti

Torp-Donner, H. ^& Juga, J.Sustainability^- animalproduction and breeding et al. 1993).The focus should be on merging

ecology and economicsso astoensure that what is economically sound in the shortterm is_eco- logically sound in the longterm (Heitschmidtet al. 1996).

In the Finnish agri-environmental programme the^costsof environmental protectionareevalu- ated and premiums arepaid _tofarmers accord- ing_tothose evaluations. The premiums for live- stock production thatarepaid^atthe^momentare for: organic production, riparian^zones,treatment of run-offwatersfrom arableland,balanced_use ofmanure nutrients, landscape and biodiversity management, extensive production and produc- tion based on local breeds(TheMinistry of Ag- riculture and Forestry 1995).

Costs ^of sustainable breeding strategies

The conservation of genetic material generates extra costs,but the possible ^returnsmay also be large. Unfortunately the_returns arehardtoquan- tify because future needs and conditions_cannot be predicted. Some principles in conservation are: ^to store small samples of manystocks, ^to choose diversestocks, to store stocks with spe- cial traits and to storelocally adapted breeds (Smith 1984). However,continuous genetic im- provement incurrentstocks may make it increas- ingly difficult for unimproved conserved stocks tocompete, unless therearereversals in breed- ing goals ordrastic changes in husbandry prac- tices (Smith ^1984).The expected benefit ⁽B⁾ in any year might be expressed by the equation: B

= P (R ^- R0) ^- nC, where P is the probability thatoneof the conserved stocks has aperform- ance greater than the original stock and so has an economic returnR which is higher than the return R()from the original stocks,and n stocks are stored, each at cost C (Smith 1984). With many stocksstored,the probability of gettingone stock with better performance than the original is increased.However,the above equation does not include the^costsof introgression and test- ing thenewgenesoranimals. Thosecostsmight become substantial especially ifthere is onlyone

gene wanted from the conserved stock, which also could be probable^(Groen and Smith 1995).

The costs of gene transfer and testing are at present high, sothe benefit should also be very large(Smith 1984).

Very long-termsupportprogrammes, partic- ularlyathigh discountrates,require an enormous ultimate pay off if thecostsare tobe recovered (Cunningham 1995).However, the normal hori- zonfor such programmes is similarto ahuman generation, and discountrates of under 5%are commonly applied in _caseslike this. As arough guide,_onecould say thatanexpected benefit of 100-200 times the annualcostcould repaya50- year investmentin the conservation ofabreed (Cunningham 1995).The probability ofabene- ficial characteristic being found in the future from_agene bank population is very small.

ndicators of sustainability

Proposals ^for implementation

Heinonen(1995)proposed modelling for sustain- able agriculture in an index form. This index aims ^to combine all theaspects that influence sustainability of agriculturesystemsand contains eight mainindicators, all of which containsev- eral variables. It is builttohave values from0^to

10, where 10 describesawholly sustainable pro- duction system. The indicators are: I) Human (the enjoyability ofwork); 2) Dependence on outside energy(fertilisers, etc.); 3)Environmen- taleffects; 4)Economics; 5) Biological efficien- cy; 6) Animal (need, etc.); 7) Social aspects; 8) Soil (erosion, etc.) (Heinonen 1995).This kind of model cannot be used in divergent produc- tion systems as the principles are quite general, but the main idea could be applied^todivergent livestock production systemsalthough more re- search for the modelling is needed.

In another research paper about the possibil- ities of modelling sustainability, the criteriawere derived from explicit but complex issues ofun-

sustainability(deWitet al, 1995),which result- ed in the conclusion thatasystem-specific anal- ysis is neededtoassessthe overall effect oflive- stock inclusion inanagricultural system oneach of the proposed general criteria for sustainabili- ty.The criteria of unsustainability in thatresearch weredefined_as: land scarcity, soil degradation, inefficient_useofresources,environmental deg- radation and declining biodiversity(deWitetal.

1995).The criteriawerequite general, and from these criteria it isalong waytodeveloping prac- ticalmeasurementsof sustainability of livestock production.

Discussion

The environmental effects of breeding pro- grammesareofgreatimportance for future live- stock production. If food production is to be maintainedat thepresent level, it is unwise to decrease the intensity of production, when the increased number of animals would thus produce morenegative environmental effects. At least at intermediate levels, more intense production decreases degradation. However, it is not clear how much of the degradation, suchas methane emissions,does notrecycle naturally into grass and crop production for livestock food. Nor is it obvious that increased intensification will reduce environmental problemsatvery intense levels.

The interactions between divergent environ- ments, management systems and intensity lev- els and production traits should be evaluatedto find_a sustainable way of animal production.

The traits that affect the longevity ofanimals, like viability,health, genetic resistance and fer- tility, have lower heritability than do production traits,and thus genetic progress in those traits is

not as easy to attain with traditional breeding methods.However, these traits are ofgreatim- portance in ethical livestock production and their impact on economics is often ignored. In big European livestock production countries, these traits _are not included in the breeding goalsas they_are in Nordic countries. Alternative breed- ing programmes (e.g. MOET), the use of new techniques like marker-assisted selection and biotechnological methods like gene transfer should bestudied,todiscover how they enhance achievements in breeding, especially for traits that have low heritability, and how they canin- creasethe competitiveness of differentbreeding populations and hence maintain genetic diversi- ty.Although it is not clear thatconsumers will acceptgenetic manipulation in food products, the ethical aspect should be studied all the same time.

Genetically variable populations are more capable of adapting to in new situations than populations that have been developed for very narrow breeding goals. Therefore the genetic diversity of production animals should be eval- uatedon a national basis _as wellas on a global basis. The breeding goals for each breed should be evaluated considering the special traits that should be maintained and used in production.

Also the effects of breeding programmes^onge- netic diversity should be evaluated both nation- ally and globally. The possible costsand bene- fits due togene conservation programmes and introgression should be taken in account when planning breeding and conservation pro- grammes.Finally, acriteriaon which breedcon- servation is based should be laid down.

Acknowledgements.WearegratefultoEsaMäntysaari,the editor ofAgriculturaland Food Sciencein Finland, profes- sorKalleMaijala,the staff at theAgricultural Universityof Norwayand two referees for their useful comments.

Torp-Donner, H. ^&Juga, J. Sustainability^- animal production and breeding

References

Beilharz, R., Luxford,B. ^&Wilkinson J. 1993.Quantita- tive genetics and evolution: ^Isourunderstandingof geneticssufficient to explain evolution? Journal of Animal Breeding and Genetics 110/3: 161-170.

Crow, J.F.1986. Basic conceptsinpopulation, quantita- tive, and evolutionary genetics.W.H. Freeman and company, New York.273p.

Cunningham,P. 1995.^Genetics,originand conservation of domestic bovids. 35p. (Mimeogr. available at De- partment of Genetics, Trinity College, Dublin 2,Ire- land).

Dempfle,L. 1990.Conservation,creation,and utilization ofgenetic variation. Journal of Animal Science73:

2593-2600.

ElTill,^A., Boiler,E.F. ^&Gendrier, J.P. 1993.Integrated Production. Principles ^and Technical Guidelines.

lOBC/WPFISBulletin16,1: 1-38.

Falconer, D.S. 1989. Introduction to quantitative genet- ics. Fourth edition,Longman ^Scientific&Technical, USA.464p.

Flachovsky, G. 1994. SindunsereMilchkuhe “Umwelts- under"? Milchrind. Journal fur Zuchtung, Biotech- nologieund Leistungsprufung^{4/94: 12-15.}

Gibson,^J.P.1994.Short-term gain attheexpenceoflong term response with selection of identified loci. In:

Proceedingsof the sth WGALP. Vol.21.^p.201-204.

Goddard, M.^1992.Optimaleffective population size for the global population of^{black and}white dairy cattle.

Journal of Dairy Science 75:2902-291.

Groen,A.F.^& Smith, C. 1995. A stochastic simulation studyof the efficiency of marker-assisted introgres- sion in livestock. Journal of Animal Breeding and Genetics 12/95: 161-170.

Haila, Y. 1995. Biodiversiteetti ja luonnonsuojelu.In:

Hiedanpää, J.(ed.) Biodiversityand production.

Satakunnan ympäristöntutkimuskeskus^{&Porin kou-} lutus-jatutkimuskeskus,p. 27- 40.

Haltia,S. ^& Mäntysaari, E, 1993.Eläinmalli^on tulossa kantakoeindeksien laskentaan. Sika 4/95:5.

Heinonen, E. 1995. Kestävän kehityksen ^mukainen maataloustuotanto Suomessa. Maa-ja Metsätalous- ministeriö,Luonnonvaraneuvosto. MMM.njulkaisuja 3/1995. 54p.

Heitschmidt, R.K.,Short, R E.^& Grings,E.E. 1996.Eco- systems, sustainability, and animal agriculture. Jour-

nal of^AnimalScience74: 1395-1404.

Hermans, C.M.L. ^&Vereijken, P.H. 1994.Grazing^hus- bandrybased on sustainable nutrient management.

In:Biologicalbasis of sustainable animal production.

EAAPPublication^No.67. ^p. 113-122.

Hill, W. 1989. Mutation and maintenance ofquantitative geneticvariation.In: Hill, W.^&Mackay, T.F.C. (eds.).

Evolution and Animal Breeding. Reviews onmolecu- lar and quantitative approaches in honour of Alan Robertson.C.A.B. International, p. 105-111.

Hodges, J. 1991. Sustainabledevelopmentof animal geneticresources. World review of animal zootech-

nie. Animal Genetic Resources 3/91.^p.2-10.

Kimura, M. 1989.Neutral theory.In: Hill, W. ^&Mackay, T.F.C. (eds.).Evolution and animalbreeding. Reviews onmolecular and quantitative approachesinhonour of^AlanRobertson. C.A.B. International,p. 13-16.

Korhonen, T.^1996. Tuotosjalostustukee kestävää kehi- tystä. Nauta 3/96. p.26-27.

Leensfra,F.R.^&Ehlhardt,D.A. 1994.Breeding goals^for intensive^butsustainable poultry meat production.In:

Biologicalbasis of sustainable animal production.

EAAPPublication No. 67.p. 169-175.

Lindhé, B. 1995.Volvo-Qualitätin Skandinaviens Stäl- len. Tierzuchter A/95. p.30-33.

Mackay,T. 1989.Mutation and the origin of quantitative variation. In: Hill, W.^& Mackay,F. (eds.). Evolution and Animal Breeding. ^Reviews on molecular and quantitative approachesin honour of Alan Robert- son. C.A.B. International,p. ^111-119.

Matassino, D., Zucchi,G. ^& diBerardino, D. 1991.Man- agement of consumption,demand, supplyand ex- changes. EAAPPublication No.48: 105-126.

Meuwissen, J.^&Woolliams, J. 1994a.Effective sizes of livestock populations to preventadeclineinfitness.

Theoretical^andApplied Genetics^{89; 1019-1026,}

- & Woolliams, ^J, 1994b. Response versus risk in

breeding schemes. Proceedings of the sth World CongressonGenetics Applied toLivestock Produc- tion. Guelph, Ontario, Canada. Vol. 18,p.^236-243.

The Ministry of Agriculture and Forestry 1995.The agri- environmental programme for Finland. MMM 17.2.1995,updatedin 20.10.1995.42p.

- 1996. Uusiutuvat luonnonvaratja biologinenmonimuo- toisuus. MMM:n biodiversiteettityöryhmänehdotus monimuotoisuuden kestävästä käytöstä. Työryhmä- muistio 1/1996.79p.

Nylander,A.^&Mäntysaari,E. 1991.Valtakunnalliset ar- vosteluteläinmallilla. Sika 2/91. p.B.

Oltjen,W.^&Beckett,L. 1996. Role of ruminant livestock insustainable agricultural systems. Journal ofAni- mal Science74: 1406-1409.

Pösö, J.^&Mäntysaari,E. 1996a.Relationships^between clinical mastitis, ^somaticcell score,^and production forthe first three lactations of FinnishAyrshire.Jour- nal of Dairy Science79: 1284- 1291.

- & Mäntysaari, E. 1996b. Genetic relationships be-

tween reproductive^disorders,operational days open andmilkyield.Livestock Production Science^{79: 41-} 48.

Puonti, M.^& Schulman,A. 1988, Eradication of haloth- ane gene from Finnish pig population. In: VIWorld conferenceon animalproduction,^Helsinki,p. 478.

Sihvonen,T. 1993. KotieläintuotteidenJa-tuotannoneet- tisyystaloudellisena tekijänä. Maatalouspolitiikan pro gradu-työ. Helsingin yliopiston taloustieteen laitos.

80p.

Smith, C. ^1984.Genetic aspectsof conservationinfarm livestock. Livestock Production Science 11: 37-48.

Vavra, M. 1996.Sustainabilityof animal production sys- tems:an ecological perspective. Journal of Animal

Science74: 1418-1423.

Wit, ^{J, de,}Oldenbrock, J.K.,vanKeulen, H.^&Zwart, D.

1995.Criteriafor sustainable livestock production;a proposalfor implementation. Agriculture, Ecosystems

&Environment53: 219-229.

Yläranta, T. 1991. Maataloustuotannon vaikutus kasvi- huoneilmiöön Suomessa. Kasvihuonekaasu- päästöjen vähentäminen. Maatalouden tut- kimuskeskus, Tiedote5/1991. 18p.

SELOSTUS

Kestävän kehityksen vaatimukset kotieläintuotannossa ja -jalostuksessa

HeidiTorp-Donner jaJarmoJuga Helsingin yliopisto

Kestävän maataloustuotannon tavoitteenaonkäyttää luonnonvaroja taloudellisesti, yhteiskunnallisesti ja ekologisestikestävällä tavalla. Kestäväkäyttö ei ku- lutapääomaa, vaan luonnonvarojen uusiutumiskyky säilyy janiiden^määräpysyy vähintään entisen suu- ruisena, Luonnonelinkykyä ylläpitää myös lajien si-

säinenjaniiden välinenmonimuotoisuus eligeneet- tinen diversiteetti. Biodiversiteetin säilyttäminen ta- kaa myös ekosysteemien monimuotoisuuden,mikä edellyttää monenlaistenkasvu-ja elinpaikkojen ole- massaoloa. Tämäkirjallisuuskatsaus perehtyy koti- eläintuotannon jakotieläinten jalostuksen vaikutuk- siinjamahdollisuuksiin kestävän kehityksen turvaa- misessa. Lisätutkimusta tarvitaanoptimoitujen jalos- tusohjelmien löytämiseksi, sillä tuotannon intensitee- tinnousu/eläinyksikkökeskinkertaisilla tuotostasoilla vähentää ympäristöhaittoja,mutta tuotantointensisee-

tinedelleen noustessaei ympäristöhaittojen vähene- minenenää olekaan selvää. Oikeapainotus kestävyy- den, eettisten arvojen, taloudellisuuden ja tuotanto-

ominaisuuksien välilläonlöydettävä.

Kaikkia jalostuskohteitaei biodiversiteetin takia kannata ottaayhteen jalostusohjelmaan, vaan ^eri- koisominaisuuksien säilyttämisen taloudellinen arvo eri roduilla onarvioitava. Kestävyyden, tuotantosai- rauksien ja geneettisenresistenssin periytymisasteet tuleemäärittää,jottavoidaan löytää edistymisenkan- nalta tarkoituksenmukaisetpainotukset. Lisäksi uu-

sientekniikoiden,kutenalkionsiirron,geenisiirron ja markkeriavusteisen valinnankäyttöä jalostuksenedis- täjinä ja monimuotoisuudenylläpitäjinä varsinkin heikosti periytyvissä terveys-,hedelmällisyys- jare-

sistenssiominaisuuksissa on mahdollisuuksien mu- kaan tehostettava.

Eri ominaisuuksien vuorovaikutukset eri ruokin- tamuotojen, tuotantoympäristöjen ja olosuhteiden kanssa vaihtelevatja järkevä yhdistelmä saattaaolla hyvinkin erilainen eri maissa. Juuri tähän erilaistu-

miseenpohjautuu jalostuspopulaatioiden maailman- laajuinen vaihtelu. Kukintuotantoeläinlaji ja-systee- mion mallitettava eripääkohdat huomioon ottaen.

Geenipankkitoiminnan ja säilytettyjen geenien käyt- töönotontodellisettaloudelliset kustannukset ja hyö- dyt vaihtelevat suuresti säilytettävänrodun erikois- ominaisuuksien taloudellistenkäyttö-jakulttuuriarvo- jen mukaan.Kansallisten jalostustavoitteiden vaiku- tusperinnölliseen muunteluun maailmanlaajuisesti juurierikoistumallaonmerkittävääja erilaisilla ja- lostusohjelmillaonmahdollisuuksiasäilyttää useampi jalostuspopulaatio kilpailukykyisenä.

Tuotannon kokonaisvaltainen mallittaminen eri aloilla tulee olemaan ehto kestävälle kehitykselle.

Jalostuksen sopeuttaminenkestävän kehityksen vaa- timuksiin ei ole ristiriidassa nykyisen linjan kanssa, vaankyse onjärkevästitoteutettavastaoptimointipro- sessista.