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Published 2003

1 History of Silage

J. M. WILKINSON
School of Biology
University of Leeds, United Kingdom

K.K.BOLSEN
Department of Animal Sciences and Industry
Kansas State University, Manhattan, Kansas

C.J.LIN
The Mennel Milling Co.
Roanoke, Virginia

What is silage? Literally, silage is anything stored in a silo, which may comprise a
hole in the ground, a bunker, a tower, a covered heap, or a wrapped bale. The word
"silage" derives from the Greek "siros" meaning a pit or a hole sunk in the ground
for storing corn (Zea mays L.; McDonald et aI., 1991). Missile weapons and dry
grain are stored in silos, but neither is described as silage-the term is reserved for
the fermented products of agricultural crops.
Woolford (1984) defined silage as "the product formed when grass or other
material of sufficiently high moisture content, liable to spoilage by aerobic mi-
croorganisms, is stored anaerobically". Silage is produced by ensilage, that is, the
placing of crop material inside a vessel or a structure called a silo. The material may
be an entire crop or only part of a crop, such as the grain portion. By placing the
material inside a structure the objective is to preserve it and to prevent it rotting as
in a compost heap. The crucial difference between a silo and a compost heap is that
air moves relatively freely in the compost heap but not in the silo.
Silage is often compared with hay made from the same crop, and the most
obvious differences are in the concentration of water and in the pH (acidity) of the
two materials. The ensiling process is potentially as efficient as haymaking with
regard to the preservation of the important nutrients in forage crops (see, for ex-
ample, reviews by Ekern et aI., 1975; Demarquilly & Dulphy, 1977; Waldo, 1977;
Zimmer, 1977, 1980). Haymaking involves the removal of most of the water in the
crop so that spoilage microorganisms do not develop during the storage period when
the crop is stored in the presence of air and not in a sealed silo. Thus, in reality, hay
crops can suffer from significant damage during the period of field drying if weather
conditions are not favorable, with loss of leaf and reduced protein content, espe-
cially in the case of legume crops like alfalfa (Medicago sativa L.; Thomas et aI.,

Copyright © 2003. American Society of Agronomy, Crop Science Society of America, Soil Science So-
ciety of America, 677 S. Segoe Rd. , Madison, WI 53711 , USA. Silage Science and Technologv. Agron-
omy Monograph no. 42.

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2 WILKINSON ET AL.

Table I-I. Composition of silage and hay made from the same original crop of alfalfa (Thomas et aI.,
1968).
Hay Silage

Water, g kg- 1 fresh


weight 148 734
Dry matter, g kg- 1 fresh weight 852 266
Ash, g kg- 1 DM 64 79
Gross energy, MJ kg- 1 DM 18.7 19.7
Crude protein, g kg- 1 DM 160 174
Crude fiber, g kg- 1 DM 325 343
NDF, g kg- 1 DM 524 495
ADF, g kg- 1 DM 381 417
Hemicellulose (g kg- 1 DM) 144 77
Lignin (g kg- 1 DM) 70 85
pH ND 4.4
Lactic acid, g kg- 1 DM NO 23
Digestibility ofDM, g kg- 1 (sheep) 607 553
Digestibility of ADF, g kg- 1 (sheep) 589 450
Digestibility ofNOF, g kg- 1 (sheep) 492 415
DM intake, g kg- 1 LW (sheep) 31.1 28.5
DM intake, g kg- 1 LW (cattle) 22.6 17.0
ND =Not determined.

1968; BoIsen, 1985) and a decrease in the digestibility ofthe organic matter (De-
marquilly & Jarrige, 1970; BoIsen et al. 1991). Hay also can deteriorate significantly
if exposed to rain or snow during storage, so a covered store or barn usually is pro-
vided to protect hay from weather damage.
The composition and nutritional value of hay and silage made from the same
original crop are shown in Table 1-1. The main differences in this particular com-
parison are the higher concentrations of water, fiber, and gross energy in the silage
than in the hay and lower values for digestibility and voluntary feed intake for the
silage than for the hay, possibly reflecting greater losses in storage for the silage
than for the hay (Thomas et aI., 1968). In this work, conducted over 5 yr and in-
volving six different crops, the results obtained in different years and with differ-
ent harvests in the same year were variable. The authors concluded that the di-
gestibility and protein values were generally higher for silages than for comparable
hays and that during the six feeding trials more dry matter was consumed as hay
than as silage, yet, because of lower digestibility, animal performance was not con-
sistently greater for hay.
Three important factors are necessary for proper ensiling-(i) agricultural crop
material, (ii) moisture, and (iii) the exhaustion of oxygen inside the silo. A silo is
the vessel or structure which is sealed to prevent the movement of air into and out
of the crop mass. The process of ensilage involves acidification of the crop by the
products of the fermentation of sugars within the plant material. The fermentation
products are organic acids, principally lactic acid. The fermentation process is sim-
ilar to that which occurs in the production of yogurt and cheese (Fung, 1988) and
in the preservation of cabbage (Brassica oleracea L.) as sauerkraut (Woolford,
1984). The acidification is the natural result of the metabolism of bacteria present
on the crop at the time of harvest. The types of acid and their amounts depend on
the amount of moisture in the crop at the time of ensiling and on the relative pop-
HISTORY 3

ulations of the different species of bacteria on the crop, known as epiphytic mi-
croflora.
Adequate acidification is vital to successful preservation of the crop, espe-
cially when the moisture concentration of the crop is relatively high, because the
acidity prevents the development of spoilage microorganisms, which are less tol-
erant of acid conditions than the lactic acid bacteria (Woolford, 1984; McDonald
et aI., 1991).
Therefore, silage is defined as an acidic, fermented, stored product from an
agricultural crop.

ORIGINS OF SILAGE MAKING

Silage making is probably more than 3000 yr old. The ancient Egyptians and
Greeks stored grain and whole forage crops in silos. Reviews of the history of silage
refer to the mural in the Naples Museum, which shows whole-crop cereals being
harvested and loaded into a small stone-built silo (Shukking, 1976; Woolford,
1984). Kirstein (1963) mentioned that silos were found in the ruins of Carthage,
indicating that forage was ensiled there at around 1200 Be. Cato (cited by Shukking,
1976) noted that the Teutons in the first century stored green fodder in pits in the
ground and then covered the pits with dung.
Little is known about silage making between about AD 100 and the eighteenth
century, although it is likely that ensiling of forage crops continued to be practiced
on a small scale throughout this period. According to Shukking (1976), grass was
ensiled in Italy in the thirteenth century, and ensiling was practiced in the northern
Alps, Sweden, and the Baltic region early in the eighteenth century. By the middle
of the nineteenth century, interest in the ensiling of grass, sugarbeet tops (Beta vul-
garis L.), and other crops had spread beyond the Baltic and Germany to most other
European countries.
In a recent, detailed, historical review of silage making in Britain, Brassley
(1996) emphasized the link between the ensiling of agricultural crops and the mak-
ing of sauerkraut, a technique that had been practiced in Germany for centuries prior
to the nineteenth. Possibly the increased movement of scientists and of scientific
information around Europe and North America led to the growth of interest in en-
siling in the nineteenth century. For example, Brassley (1996) cited an article in the
Transactions of the Highland Agricultural Society in 1843, in which James John-
ston, a lecturer in Chemistry at the University of Durham, posed the question: "Is
it possible to preserve crops in their moist state?" He then translated an article by
Grieswald (also cited by Watson, 1939) in the Transactions of the Baltic Associa-
tionfor the Advancement of Agriculture for 1842 that described the preservation
of grass by salting-the same technique used in the making of sauerkraut. Brass-
ley argued that the ensiling of crops in the nineteenth century probably was devel-
oped in northern Europe, in the Baltic region or in Germany.
From Germany, the focus of the development of ensiling in the second half
of the nineteenth century moved to France, inspired by the publication in France
of letters to a German newspaper by a farmer from Stuttgart, Herr Reichlen, who
had experimented with the ensiling of sugarbeet leaves and tops and of whole-crop
com in long, narrow trench silos. The letters were translated into French and pub-
4 WILKINSON ET AL.

Ii shed in the Journal d'Agriculture Practique in 1870. Then, in 1877, the French
farmer Auguste Goffart published the results of his work on the ensiling of chopped
forage com and other crops (Goffart, 1877). Goffart's book was influential not only
in Europe but also in North America, where it was published in 1879 by J.B.
Brown, the President of the New York Plow Co., who sent copies to his customers
as an advertisement for his company (Carrier, 1920). However, Brassley (1996)
stated that the first silo in the USA was probably built in 1873 in Illinois by a farmer
who had read an English translation of the articles in the Journal d'Agriculture Prac-
tique while he was a student at the University of Illinois. Influenced by the French
reports, M. Miles, a farmer in Lansing, MI, experimented with ensiling whole-plant
com in 1875. In an article published in Country Gentleman on 5 Oct. 1876, Miles
described his own silage making experiences and those in France. Miles used the
word "silo", and he suggested the adoption of "ensilage", in the absence of an Eng-
lish equivalent for the fermented product (Miles, 1910). By 1882, a report by the
United States Department of Agriculture contained statements from more than 90
farmers in both the USA and Canada regarding their ensilage practices. Goffart
(1877) described the chopping of the whole com plant into lO-mm lengths and en-
siling the chopped material in large bins built of brick (4 m wide by 10m long, and
5 m high). These silos probably were considered to be better suited to the small farms
in France than Reichlen's trench silos. Nevertheless, it is most likely that Reichlen,
not Goffart, was the innovator responsible for the intense interest in silage making
that developed towards the end of the nineteenth century on both sides of the At-
lantic.
At the same time, Samuel Jonas, a farmer in Britain, published details of his
technique for ensiling chaff by mixing it with 50 kg of green rye (Secale cereale
L.) and "one bushel" (36.3 kg) of salt per tonne of chaffin the Journal of the Royal
Agricultural Society. He described how the silage had enabled him to feed his sheep
(Ovis aries) and cattle (Bos taurus) during two winters when his turnip (Brassica
rapa L.) crop (a traditional winter livestock feed along with hay) had failed com-
pletely (Jonas, 1870). In 1874, the Professor of Agriculture at Royal Agricultural
College undertook a tour of the Austro-Hungarian Empire and in his report described
the production of "sour hay" "by digging long graves or trenches, 4 feet by 6 or 8
feet in depth and breadth and cramming the green grass or green Indian com tightly
down into them, covering the whole with one foot of earth .... no salt is used and the
operation is as simple as it appears in the description" (Wrightson, 1874).
Several leading British farmers began experimenting with silage making, in-
spired by visits to other countries such as Holland, where Goffart's book had been
published in Dutch (Shukking, 1976). Leading farmers from France visited agri-
cultural shows in England to demonstrate their techniques. One farmer in Wales, a
Mr. Kenyon, made silage in 1881 in a brick silo with a concrete base and a drain,
which led to a stream. The silo had a lid which was able to move down the per-
pendicular walls as the contents subsided after filling. The silo also had a roof to
keep out rain. The grass crops were chopped to 25-mm average lengths by a chaff
cutter powered by water. Mr. Kenyon apparently learned how to make silage in air-
tight conditions while living in the USA before he moved to Wales. His knowledge
and success probably predated the French influence on English farmers (Jones,
1992).
HISTORY 5

In 1883, the British Government instigated the first official study of silage
making by asking the Royal Agricultural Society to commission a survey of silage
making in the British Isles and elsewhere. The results of the survey, which involved
36 farms, were published in 1884 (Jenkins, 1884). For a summary of the survey,
see Brassley (1996), who mentioned some of the more significant findings, including
the first report of the pollution of drinking water by silage effluent, the first report
of portable silos (wooden "bales"), the first report in Europe of blowers for filling
silos, and the first report of the feeding of silage effluent to pigs (Sus scrofa). The
conclusions of the survey were that corn was the best crop for silage, that the feed-
ing value of silage was similar to that of hay, but that silage was not a complete sub-
stitute for hay. The Society's chemist reported on the analysis of samples of the silage
made on farms involved in the survey (Voelcker, 1884). He concluded that the en-
siling process was a bacterial process and distinguished between "sweet" silage,
which did not have a strong odor, and "sour" silage, which did and presumably con-
tained acetic acid and/or butyric acid.
Meanwhile, Goffart's work was taken up enthusiastically in the USA . Rew
(1888) described the delighted reactions of one Vermont farmer to the making of
silage from whole-plant corn at the Fifth Ensilage Congress in New York in 1886.
In 1889, Miles published his first book, entitled Silos, Ensilage, and Silage, A Prac-
tical Treatise on the Ensilage of Fodder Corn. By the end of the nineteenth century,
most Land Grant universities were making silage and promoting its use on farms .
Several universities also were conducting silage research (Hunter & Bushnell,
1916; Reed & Fitch, 1917; Eckles et aI., 1919; Hulbert & Christ, 1926). Kansas State
Agriculture College published Silos and Silage (Bulletin No. 6) in 1889, which re-
ported cattle performance and the sources of loss in an 80-t capacity tower silo. Seven
percent of the weight of the whole-plant corn ensiled vs. weight of silage removed
could not be accounted for, so the authors explained it as a loss by "evaporation".
The initial enthusiasm for silage in the UK was reflected in the encourage-
ment for the adoption of the technology from government, agricultural societies,
and leading landowners, who recognized that the main advantage of silage was that
it was less susceptible to total failure than either hay or root crops. Unfortunately,
the adoption of silage making in the UK and elsewhere in Europe suffered a severe
setback with the publication of the book Sweet Ensilage (Fry, 1885). While Fry rec-
ognized the importance of having an air-tight silo, he advocated that all crops must
reach a temperature of at least 50°C before the silo was sealed. In Germany, Kell-
ner (1905) considered acid silage preferable to sweet ensilage, but that the technique
of ensiling was useful only when other methods were likely to lead to a complete
loss of feed.
The advocacy by Fry (1885) of delaying the sealing of silos until the crop had
heated to 50°C or more persisted in some parts of Europe until the early I 960s.
Moore (1950) states that too Iowa temperature could be a worse fault than too high
a temperature, and that at high temperatures (>50°C), "no undesirable fermenta-
tion is likely to take place". The "sweet", dark brown product undoubtedly under-
went a caramelization of plant sugars and Maillard complexing of sugars with pro-
teins (McDonald et aI., 1991). Moore recognized that although sweet silage was
palatable to livestock, the digestibility of the protein was substantially reduced. It
was considered that the lactic acid organisms grew best at high temperatures, and
6 WILKINSON ET AL.

that if the silage was allowed to heat up, the risk of producing a foul-smelling bu-
tyric acid fermentation was reduced. Consequently, farmers equipped themselves
with thermometers that were placed in the forage mass in the silo to ensure that the
material heated up adequately. Moore recalled recording the temperature of ensiled
material in an unwalled clamp silo on a farm in Staffordshire, UK, for three con-
secutive days in 1960 after the harvest had been completed and the ensiled mate-
rial rolled. The silo was not sealed, and the silage produced was black and smelled
of rotted compost on the outside and of tobacco (Nicotiana tabacum L.) in the cen-
ter. Not surprisingly, the material was virtually inedible when offered to dairy
cows.
The concept of sweet silage was mistaken and counter-productive. The no-
tion that silos could be built on a small scale without walls, with delayed and in-
adequate sealing of silos with earth or manure in areas of high rainfall and with lit-
tle chopping of the crop at harvest all combined to produce overheated material,
high losses of nutrients due to excessive oxidation, poor preservation quality, and
low feeding value. Not surprisingly, silage making failed to develop for many
decades from its initial promising origins. For a fuller discussion of these and other
issues that contributed to the delayed adoption of silage making, see Brassley
(1996).

MILESTONES IN SILAGE MAKING

The production of so-called sweet silage remained a millstone around the neck
of good silage making practice in some regions of Europe for the first half of the
twentieth century. Nevertheless, silage making technology advanced steadily in other
parts of the world. The development of tower silos was probably the first major mile-
stone in the successful development of silage making in Europe and North Amer-
ica. Schukking (1976) described the Cremasco tower silos in Italy, which were pop-
ular at the beginning of the twentieth century. The technique involved ensiling wilted
crops of 50 to 70% dry matter in air-tight concrete upright silos. By the 1920s, sev-
eral thousand Cremasco silos were erected in the region of the Po valley, where Ital-
ian ryegrass (Lolium multiflorum L.) was the major forage crop, grown with fertil-
izer N and irrigation.
Tower silos proved popular in many other areas of Europe and in North Amer-
ica. Many small tower silos were built in the 1920s from wood, tile, or concrete-
some sited inside buildings to protect the silage from freezing solid in winter and
to provide good accessibility of the silage to livestock. Some of these early silos
have survived to this day. The number of tower silos erected on farms in North Amer-
ica increased from about 2000 between 1880 and 1900 to more than 100 000 by
1925. As recently as 1974, about 4000 tower silos were built annually, but the num-
ber of new tower silos erected in the 1980s and 1990s has declined dramatically.
As farms expanded in size and livestock numbers increased, more and more farm-
ers selected horizontal storage systems, such as bunkers, trenches, drive-over piles,
and bags, to meet their additional silage requirements. However, recent innovations
in the design and engineering of silos and silage un10aders make it possible to store
nearly 2000 t of dry matter in one tower silo and unload up to 150 kg dry matter
HISTORY 7

min- 1 (D. Freeman, Crop Storage Institute, Lafayette, IN, personal communication,
1997).
The advantages of the tower silo over all other early types of silos were that
it had an airtight base, airtight sides, and a relatively small exposed area at the top,
which mayor may not have been sealed. The dimensions of the tower favored com-
paction of unchopped material over bunker, trench, or pile silos. With no sealing
of the sides and the unchopped grass layered horizontally, air ingress into the mass
of material in the silo was severe and oxidative losses were very high as a result.
Sweet silage was so popular and the fear of cool fermentations in wet crops
giving rise to poor quality material so great that some early silage makers actually
advocated heating the silos or the crops at ensiling to ensure that the temperature
of the crop within the silo was raised to 50°C. The processes, described by Watson
and Nash (1960), included heating the crop with steam, immersing the crop in a
water bath kept at 100°C immediately prior to ensiling, sterilizing the crop with pres-
surized steam prior to ensiling, heating tower silos or pits by electrically heated el-
ements (known in the 1920s as "electro-silage"), and heating air electrically and
forcing it through a drain pipe within the silo. All these methods were found to have
little effect on the quality of silage and simply added to the cost of the ensiling tech-
nology and, consequently, were soon abandoned.
Farmers have been using various additives or "supplements" throughout the
twentieth century in the hope of making better silage (McCullough, 1977). In the
early years of silage production, the reason for applying an additive was to prevent
secondary fermentation and a butyric acid silage. As a result, the efficacy of the ad-
ditive was judged by its effect on typical fermentation criteria, such as pH, and con-
tents of ammonia-N and lactic, acetic, and butyric acids (Spoelstra, 1991). This ori-
entation on fermentation was reflected in the traditional division of additives into
categories of fermentation inhibitors, fermentation stimulants, and substrate or nu-
trient sources (Woolford, 1984; Pitt, 1990; McDonald et aI., 1991 ; Boisen, 1995).
McCullough (1978) compiled a list of several silage additive ingredients and the
earliest year of their recommended use: sodium chloride (1893), whey ( 1895), car-
bon dioxide (1900), sulfur dioxide (1910), acid salts (1912), molasses ( 1917), ben-
zoic acid (1923), and formaldehyde (1931).
An early system used to judge the quality of silage was the Flieg point
scheme, which awarded points according to the relative amounts of lactic, acetic ,
and butyric acids: the higher the proportions of lactic and acetic acids to butyric acid,
the higher the score and the better the quality (Flieg, 1938). This scheme gained
wide acceptance by farmers in Europe, especially in Germany. Later, Zimmer
(1966) modified the Flieg point scheme, and less emphasis was placed on the con-
tent of butyric acid, presumably in response to the greater incidence of well-pre-
served silages on farms.
A milestone of great significance was reached with the publication of work
on the direct acidification of crops at ensiling in Italy (Giglioli, 1914), in Germany
(by Fingerling) who both advocated the use of dilute hydrochloric acid, and by A.I.
Virtanen (in Finland) who worked mainly with a mixture of hydrochloric and sul-
phuric acids. These acids were added to the crop manually by use of watering cans,
which made the process laborious and hazardous. Other inorganic and organic acids
were investigated in the 1920s and 1930s along with salts of acids. For detailed re-
8 WILKINSON ET AL.

Table 1-2. Effects of direct acidification of clover (Trifolium spp.) with hydrochloric acid on protein
breakdown.t
HCI added

Item Fresh crop High Medium Low No acid

Initial pH 3.7 4.1 4.5


Final pH 3.6 4.3 4.6 4.5
Soluble N, g kg- 1 total N 26 28 44 60 65
NHrN, g kg- 1 total N 1.5 2 12 21 22

t Source: Virtanen, 1933, quoted by Watson & Nash, 1960.

views of the early research on additives for silage, see Watson and Nash (1960) and
Shukking (1976).
The research on the direct acidification of crops prior to ensiling was of enor-
mous and long-lasting significance for two main reasons. First, protein breakdown
was reduced, and second, the growth of harmful bacteria was prevented. Many early
silage research workers were well aware of the problems in cheese making caused
by undesirable bacteria and that the bacteria responsible might have originated in
poorly preserved silage. They were also aware that heat-damaged protein reduced
the feeding value of the silage. Virtanen (1933) summarized the effects of acidifi-
cation on the breakdown of protein and reported that if the pH of the crop could be
reduced below pH 4.0, then proteolytic enzyme activity was inhibited virtually com-
pletely (Table 1-2). The effects of acidification were summarized by Virtanen
(1933) thus: "All detrimental breakdown processes in the fodder would be elimi-
nated by treating the fodder, at the time of ensiling, with such amounts of acid as
would rapidly raise the acidity of the mass to a point below pH 4.0".
At the same time, research was proceeding in France, Germany, and North
America into the stimulation of fermentation by the addition of lactic acid bacte-
ria, molasses, and other carbohydrate sources to sugarbeet residues and other crops
(Reed & Fitch, 1917). By the 1930s, molasses was recommended for use in Ger-
many, Holland, the USA, and the UK (Watson & Nash, 1960). The concentration
of protein in the crop was considered to be the main factor likely to affect the qual-
ity of the fermentation process, and thus the amount of molasses required to ensure
efficient preservation.
Two developments in the late 1960s combined to revolutionize the use of
silage additives on farms. The first was the introduction of the gravity-feed appli-
cator, and the second was the development of formic acid as an alternative to the
direct acidification of the crop with mineral acids, which was commonly referred
to as the "AIV" process of A.1. Virtanen. These changes allowed farmers to apply
additives directly into the delivery chute or the chopping chamber of the forage har-
vester, thereby achieving uniform application of relatively low rates of additive, typ-
ically 2.5 to 4.0 L c 1 of fresh crop. Formic acid was effective because it has spe-
cific antimicrobial activity against clostridia, whereas inorganic acids simply
restricted the fermentation by reducing pH (Woolford, 1975, 1978a). However, the
volatility of formic acid was a disadvantage, and many farmers opted to use other
types of additives, such as salts of acids and bacterial inoculants, which were eas-
ier to handle and much safer for the farmer to use. If the 1970s were dominated by
HISTORY 9

the adoption by farmers in Western Europe of organic acid additives and conditioners
on mowers for grass crops, then the 1980s saw the introduction of sulphuric acid
as an additive to replace formic acid for wet crops of grass (O'Kiely et al., 1989;
Appleton, 1991; Steen, 1991).
The first known use of lactic acid bacterial (LAB) cultures was with ensiled
sugarbeet pulp in France in the 1900s (Watson & Nash, 1960). Kuchler (1926) (cited
by Spoelstra, 1991) described an inoculant system developed in Germany, which
included the growing of bacteria on the farm. However, many of the earlier attempts
to inoculate silage crops were not successful because either the strains of LAB were
not adapted to a silage environment or the bacterial cultures were not viable at the
time of use (Spoelstra, 1991). Disappointing results with some earlier inoculants,
particularly with crops low in fermentable substrate, also were reported (Haigh et
al. , 1987; Weinberg et al., 1988).
However, technology developments in the 1970s and 1980s led to improve-
ments in the commercial production of bacterial cultures used in silage inoculants.
An overview of the procedures was presented by Aimutis and Boisen (1988), and
a summary of the fermentation and stabilization techniques was reported by Ris-
ley (1992). Perhaps no other area of silage production has received as much atten-
tion among both farmers and researchers in the 1980s and I 990s as bacterial inoc-
ulants. Research led to a better understanding of the microbiology of silage (see
reviews by Lindgren, 1991; McDonald et al., 1991; Pahlow, 1991; Spoelstra, 1991).
At the same time, the crucial role of the epiphytic microflora on the crop in deter-
mining when inoculation was likely to work and when it was not reported by Muck
(1991). Inoculants became the most commonly used silage additive, particularly in
regions where com and field-wilted grasses and legumes were the major silage crops
(Boisen & Heidker, 1985; Wilkinson, 1990; Kung, 1992; Boisen et al. , 1995). It is
beyond the scope of Chapter I to cite all of the recently published scientific data
to document the effect of inoculants on silage fermentation , preservation effi-
ciency, and nutritive value. For detailed summaries on bacterial inoculants, see re-
views by Boisen and Hinds (1984); Harrison (1989), Muck and Boisen (1991),
Spoelstra (1991), Muck (1993), Castler (1990), Kung and Muck (1997), and Muck
and Kung (1997).
Although whole-plant com and sorghum [Sorghum bicolor (L.) Moench] have
always been considered easy crops to ensile, their protein concentration only ranges
from 60 to 90 g kg-I of dry matter, which is far below the requirement of most live-
stock. Research was undertaken in the I 960s, particularly at several Land Grant uni-
versities in the USA, to increase the protein equivalent by the addition of nonpro-
tein nitrogen (NPN) to the crop prior to ensiling (Owens et al. , 1969; Henderson et
al., 1970; Huber & Santana, 1972). Earlier research with urea and anhydrous am-
monia additions to com and sorghum silages was reviewed by Ely (1978). The re-
view did not include silage preservation results; however, retention of added N was
0.95 or higher for urea-treated but only 0.50 to 0.75 for ammonia-treated silages.
In a series of six trials conducted with whole-plant com and sorghum in farm-scale
silos, Boisen et al. (1992) reported that anhydrous ammonia applied at 35 to 40 g
kg-lor urea at 50 g kg-I (fresh basis) increased the lactic and acetic acid concen-
trations, but dry matter recovery was reduced in five of the six trials. Performance
of growing cattle was not improved by the NPN additions, and gain per tonne of
10 WILKINSON ET AL.

crop ensiled was reduced by an average 0 f 3.0 and 7.0 kg in the com and sorghum
silages, respectively. In a review of 39 studies published between 1985 and 1992,
Muck (1993) found that NPN additives increased the fermentation acid concen-
trations and extended bunklife but decreased dry matter recovery in a majority of
the studies, and clostridial activity was a problem in crops ensiled with less than
300 g of dry matter (DM) kg-I. Digestibility of the NPN silages (e.g., DM, NDF,
or ADF) was increased in 16 of 19 studies; however, these apparent improvements
in nutritive value did not increase daily gain, milk production, or feed efficiency in
most studies, especially in grass and legume silages. Muck noted that the apparent
paradoxes of improved aerobic stability with reduced dry matter recovery and im-
proved nutrient digestibility with no benefit in livestock performance need further
clarification.
Although anhydrous ammonia continues to be used in a few regions of North
America, it is unlikely to become a popular additive in the future unless the prob-
lems of handling, application, preservation efficiency, reduced dry matter intake,
and the increased risk of a clostridial fermentation are overcome. When economic
analysis includes the increased silage dry matter loss and the cost of replacing the
volatile N loss, ammonia can be an expensive source of supplemental protein for
beef and dairy cattle (Boisen et aI., 1998). Urea is easier and safer to handle than
ammonia, but unless future trials show significantly increased nutritive value for
the traditional silage crops (e.g., com and perennial grasses), it will not become a
commonly used additive.
The next milestone was the development of the forage harvester. It had been
known since the beginning of the twentieth century that the liberation of plant cell
sap accelerated the rate of fermentation in ensiled crops. However, not until the de-
velopment of the forage harvester in the 1940s did the technology of silage mak-
ing advance significantly beyond that of hay making. Nevertheless, the introduction
of the baler at about the same time meant that in those countries with good weather,
haymaking remained the predominant method of forage preservation. Early forage
harvesters were relatively expensive and could be afforded only on larger farms.
As a result, forage harvesters did not become popular for many years after their de-
velopment, and buckrakes and stationary green crop loaders were used, with little
or no chopping of the crop prior to ensiling.
For most of the first half of the twentieth century, crops for silage were har-
vested as for hay, by buckrake after being mowed mechanically and then raked into
swaths. The first forage harvesters were direct-cut machines; that is, they cut and
harvested the crop in the same operation. Moore (1950) stated that the main ad-
vantage of a forage harvester was a reduction by about 50% in the labor required
to make silage. Direct-cut forage harvesters were at work as early as 1950.
Following the introduction of the forage harvester, research workers soon dis-
covered that chopping the crop at the time of harvest improved the quality of the
fermentation. In a significant and influential series of experiments carried out in 1952
and 1953, Murdoch et al. (1955) found that chopping grass through a cutter-blower
into lengths of 50 to 100 mm resulted in consistently lower temperatures during the
first 14 d after ensiling, a reduction in the concentration of butyric acid in the silage,
and an increase in amino acids as a proportion of total crude protein (Table 1-3).
Balch et al. (1955) also found that the digestibility of chopped silage was higher
HISTORY 11

Table 1-3. Effect of chopping grass prior to ensiling on fermentation quality.t

Item Unchopped Chopped

Oven dry matter, g kg-I


Crop at harvest 234 230
Silage 177 197
pH 5.4 4.7
Lactic acid, g kg- 1 0M 2 26
Acetic acid, gkg- 1 0M 20 29
Propionic acid, g kg- 1 0M 9 9
Butyric acid, gkg- 1 0M 46 24
Valerie acid and higher acids, g kg- 1 0M 23 13
Volatile basest, g kg-I CP 369 199
Amino acid, g kg-I CP 188 296
t Source: Murdoch et aI. , 1955 .
:j: An indicator of protein breakdown , most likely ammonia.

than that of unchopped silage. Recently, amino acid N as a proportion of total sol-
uble N was found to be one of the three best predictors of the voluntary intake of
grass silage by dairy cows (Offer et aI., 1994).
Murdoch et al. (1955) commented on their results: "An outstanding point of
interest is the quality of the chopped silages despite the fact that they were made
at relatively low temperatures (26 to 32°C) .... This would indicate that if conditions
in the silage are suitable for bacterial growth the temperature has little importance
in the ensiling process". At last, the fallacy of hot "sweet" silage was being exposed.
The flail harvester was introduced in the USA in the early 1950s. This ma-
chine cut and loaded the grass in one single movement, by using the flails which
rotated at the same speed as impellers to drive the crop pneumatically up a chute
and into a trailer. The flail harvester was a popular machine on smaller livestock
farms for many years, and it is still in use in some areas of Europe and in several
developing countries in the tropics, where silage making was introduced in the 1980s
and I 990s.
The early work on chopping by Murdoch and others laid the foundation for
the development of improved forage harvesters, introduced in the 1960s, which were
capable of chopping or lacerating the crop as it was passed through the machine.
The pioneering English work on chopping and its effects on silage composition was
developed further in France and Belgium in detailed studies on feed intake and uti-
lization (Dulphy & Demarquilly, 1973; Dulphy et aI., 1975; Deswysen & VanbeIle,
1978). The introduction in the late 1960s and early 1970s of metered-chop forage
harvesters allowed grass to be chopped uniformly to average lengths as low as 5 to
7 mm. These machines constitute the majority of the forage harvesting equipment
in use today, though most operators now compromise between long chopping and
reduced silage quality, and very short chopping, slower harvesting, and excessive
fuel consumption.
The next major innovation to be introduced, probably around 1950, was
plastic sheeting (Shukking, 1976). The earliest sheets were made of polyvinyl
chloride and were used to protect small field clamp silos from rain ingress. By 1960,
12 WILKINSON ET AL.

sheets were increasingly made from polyethylene and included better plasticizers
and ultra-violet stabilizers to prevent the material from disintegrating in sunlight.
The sheeting was used to provide a way of separating the silage from the soil on
top. However, research in the late 1950s demonstrated clearly that without adequate
surface pressure on the top of the silo, air could penetrate the mass and severely re-
duce the quality of the silage, partly through increased temperature and greater ox-
idative losses (McDonald et aI., 1960).
Once plastic sheeting was accepted as the most practical way of sealing
bunker, trench, and pile silos, the next innovation was "vacuum" silage in the early
1960s. In this method, small piles of silage were made on a bottom sheet of plas-
tic and covered completely with a top sheet, which was held close to the bottom
sheet with soil placed around the base of the pile. A hole then was made in the top
sheet at a convenient point and a vacuum applied to suck out the air. The process
of evacuating gas from the pile was repeated until further gas evolution was negli-
gible. The technique was laborious and suffered from the fact that sustained pres-
sure was not applied on the top surface, so any holes or leaks resulted in substan-
tial surface wastage.
Not long after the introduction of vacuum silage, it was made obsolete by the
development by Richard Waltham, a Dorset farmer, of the "Dorset Wedge" system
of filling bunker, trench, and pile silos. Essentially, the method involved filling the
silo and rolling the chopped material on an inclined rather than on a horizontal plane,
which had the effect of reducing the area of crop exposed to the air and of achiev-
ing greater compaction during filling. For a full description of this important mile-
stone see Raymond and Waltham (1996).
The great benefits of plastic sheeting were that air movement into and out of
the silo could be reduced and surface waste could be minimized as a result. Yet many
years elapsed in North America before farmers began to use plastic sheets to re-
duce the surface wastage. Most silos, especially large bunkers, trenches, and piles,
were left unsealed after filling in the hope that rain or snow would provide an ef-
fective cover or protective seal for the ensiled crop. However, today a much higher
percentage of silos are being sealed with plastic sheets because the magnitude of
surface waste in unsealed silos was quantified in research by McLaughlin et al.
(1978), McGuffey and Owen (1979), Ashbell and Weinberg (1992), Dickerson et
aI. (1992), BoIsen et aI. (1993), and Holthaus et al. (1995). Computer spreadsheets
were made available that allowed farmers to estimate the value of silage saved by
sealing based on the value of the crop, silo dimensions, cost of the plastic, and cost
of the labor needed to apply and remove the sheet and weighting material (Huck et
aI., 1997). Recent data show that when top spoilage is included in a com silage-based
diet at levels as low as 125 to 250 g kg- 1 of DM, feed intake and digestibilities of
organic matter, protein, and fiber are all significantly reduced (Whitlock & BoIsen,
1998, unpublished data).
Meanwhile, large round bales were introduced in the mid 1970s along with
plastic bags, which had to be tied individually to seal the material. Unfortunately,
incomplete sealing, especially at the neck of the bag, allowed air to penetrate into
the bale and, as a result, bagged bale silages often showed evidence of mold de-
velopment and elevated levels of contamination by Listeria monocytogenes (Fen-
lon, 1988).
HISTORY 13

Stretch-wrapping of bales was introduced in the early 1980s and was adopted
rapidly by farmers who appreciated the greater efficacy of the approach and its re-
duced labor requirement. Work in Sweden (Lingvall & Nglund, 1993) showed that
wider wraps and more layers were very effective in providing a seal for grass silage.
The next milestone was the development in the early 1970s of the mower-con-
ditioner. The use of flail mowers was associated with a more rapid rate of water loss
from mown grass swaths than with the reciprocating knife mowers, which had been
used for many years for both hay and silage crops. However, flail mowers were
thought to put the crop at risk of nutrient losses through leaching, if the swath re-
ceived rain during the period of field wilting. A better system of accelerating loss
of water from mown crops was needed.
Much of the research into the mechanical treatment of grass crops at mow-
ing was carried out in Holland by Bosma and in the UK by Klinner. The subject
was reviewed by both workers (Klinner, 1976, 1984; Bosma, 1991). Conditioning
of grass crops involved passing the crop through twin rotors, on which spokes were
mounted and which revolved at different speeds. The spokes on the conditioner
abraded and punctured the cuticle of the grass, so that the evaporating area was in-
creased. Unfortunately, many of the early mower-conditioners produced a narrower,
more upright swath than mowers without conditioners, and as a result, the effects
of conditioning on rate of water loss were small. Also, conditioning was found to
have little effect if the swath was tedded soon after mowing (Bosma, 1991). Nev-
ertheless, conditioners proved to be popular additions to mowers, especially in re-
gions where tedding was not practiced (see below).
In the North America, where whole-plant com is the predominant silage crop,
forage harvesters equipped with a kernel processor have just recently caught the at-
tention of researchers, dairy farmers, and silage contractois, even though the tech-
nology was developed more than 20 yr ago and is widely used in the com silage
regions of Western Europe (Honig & Rohr, 1982). The kernel processor has helped
to minimize the risks associated with com hybrids that dry down quickly by ex-
tending the harvest window and improving the utilization of the energy and fiber
components of the silage (Bal et aI. , 1996; Harrison et aI., 1997; Young et aI., 1998).
However, for this technology to gain wide-spread acceptance, the relationships be-
tween kernel processing, com hybrid, milkline score (percentage), and chop length
need further clarification, particularly on how they affect preservation efficiency,
aerobic stability during the feedout period, and starch and fiber digestibilities (L.
Satter and R.E. Muck, Dairy Forage Research Center, Madison, WI, personal com-
munication, 1998).
Interest among farmers and researchers in alternative crops for silage increased
in the I 970s and 1980s. A few of these crops include forage sorghums (Kirch et
aI. , 1987; White, 1989), whole-crop small-grain cereals (Ashbell et aI. , 1985;
Azimi et. aI., 1988; and reviews by Oltjen & BoIsen, 1978; Wilkinson & Stark, 1990;
Stark & Wilkinson, 1992), field pea [Pisum sativum subsp. arvense (L.) Pior]
(Ashbell & Weinberg, 1991), soybean [Glycine max (L.) Merr.] (Harbers et aI.,
1992), whole-plant rice (Oryza sativa L.) (Nakui et aI., 1988), ground high mois-
ture com grain (Buchanan-Smith & Young, 1991), mutants of com hybrids with im-
proved nutrient digestibilities or amino acid composition (Cherney et aI., 1991).
These crops clearly have their place in regions where conventional crops are unre-
14 WILKINSON ET AL.

liable agronomically, or where the crop and livestock production systems are well-
suited to the inclusion of these crops. A recent milestone was the incorporation of
chopping mechanisms in big balers, so that the crop is ensiled at a higher density
than that of unchopped material. Initial results (Fychan & Jones, 1994) indicated
that chopped baled silage was likely to have a better fermentation quality and nu-
tritional value compared with unchopped silage and similar to that of silage made
by a metered-chop forage harvester and ensiled in a bunker silo. This development
is likely to accelerate the adoption of silage on smaller livestock farms that previ-
ously had encountered problems with poorly preserved silages made from relatively
wet crops into big bales.
A current major interest is improving the hygienic quality of silage, especially
with regard to reducing aerobic spoilage during the feedout period (see reviews by
Weissbach, 1996; Gotlieb, 1997; Whitlow & Hagler, 1997; Seglar, 1997). Work has
been in progress in some countries for several years, with the greatest progress being
made by German researchers (see reviews by Oldenburg, 1991; Honig, 1991 ;
Weissbach, 1996). This interest is stimulated in part by the relatively higher dry mat-
ter concentrations of silages in recent years, and also by an increasing concern that
even well-preserved silage, if it is made from contaminated crops or allowed to de-
teriorate on exposure to air during the feedout period, might be responsible for prob-
lems of livestock health and loss of productivity.

CURRENT PRODUCTION OF SILAGE AND HAY

The production of hay remained relatively similar in the UK from 1884 to


1974, and only in the last 16 yr has silage making predominated over hay making.
The decrease in the production of hay since the late 1970s probably has been the
result of the adoption by smaller farmers of big bale silage and increased use of con-
tractors (custom hire) on medium-sized farms.
In some countries of Western Europe, like Finland, silage probably has been
the major method of forage preservation for many decades (Wilkinson & Stark,
1987). By 1994, the production of silage exceeded that of hay in all 17 countries
of Western Europe except Austria, France, Greece, and Switzerland (Wilkinson et
aI., 1996). The estimated areas of the major crops harvested for silage in Europe
and North America in 1994 and estimated production of silage and hay in 1994 are
shown in Tables 1-4 and 1-5, respectively. The information was gathered from of-
ficial statistics, where available, or from estimates by individual experts. It must be
stressed that the data are estimates and, therefore, should be treated with appropri-
ate caution.
Grass, harvested from temporary and permanent pastures, is the predominant
crop for silage in Western Europe and Russia. In Eastern Europe the areas of grass
from pastures and com harvested for silage in 1994 were similar (Table 1-4).
Silage was made from significant areas of other crops in Europe, mainly sugarbeet
leaves and tops. Compared with North America, the areas of grass, com, and other
crops harvested for silage in Europe were relatively large, but the areas of legume
crops harvested for silage were relatively small. It is notable that substantial areas
of crops were harvested for silage in the Russian Federation.
HISTORY 15

Table 1-4. Estimated areas of crops harvested for silage in Europe and North America in 1994. t

Perennial Whole-crop Other


Area grasses Com Sorghum Legumes:j: cereals§ crops'll
1000 ha
Europe
W. Europe# 9646 3890 182 324 641
E. Europett 1332 1288 193 208 1577
Russia:j::j: 9736 7500 375 2251
Total 20714 12678 532 4469
North America
USA 2268 133 750
Canada 166 486 304
Total 2434 133 304 500

t Sources: Wilkinson et aI. , 1996; Wilkinson & Boisen, 1996.


:j: Mainly alfalfa.
§ Mainly wheat.
'II Mainly beet tops.
# European Union (15 countries) + Norway and Switzerland.
tt 15 countries.
:j::j: Russian Federation.

The production of hay exceeded that of silage in Eastern Europe, Russia, the
USA, and Canada. Only in Western Europe was the estimated output of silage greater
than that of hay, possibly reflecting the relative suitability of the climate for hay-
making in areas other than Western Europe. In Western Europe between 1990 and
1994, little increase has occurred in the total area of crops harvested for silage, and
a small decrease has occurred in the output of silage (Wilkinson & Stark, 1992;
Wilkinson et al., 1996). The USA has experienced a similar trend for com in the
past three or four decades-fewer hectares were harvested for silage, but there was
very little change in total output of silage. However, the amount of silage produced
in Mexico has more than doubled in the 1990s, and it is now estimated to be about
15 to 17 million tonnes offresh weight annually (G. Hoyos, Alltech, Mexico City,
Mexico, personal communication, 1997). In fresh weight terms, the total amount

Table 1-5. Estimated quantities of silage and hay made in Europe and North America in 1994. t

Area Hay Silage Total

106 t of dry matter


Europe
W. Europe:j: 60.1 91.6 151.6
E. Europe§ 32.3 15.4 47.7
Russia'll 59.7 45.0 104.7
Total 152.1 152.0 305.0
North America
USA 123.0 39.3 162.3
Canada 40.9 7.4 48.3
Total 163.9 46.7 210.6

t Source: Wilkinson & Boisen, 1996.


:j: European Union (15 countries) + Norway and Switzerland.
§ 15 countries.
:j: Russian Federation.
16 WILKINSON ET AL.

of silage made in Europe and North America in 1994 was about 550 and 130 mil-
lion tonnes, respectively.
Brazil, Argentina, and Chile in South America and Australia and New Zealand
have all experienced very rapid growth in silage production in the 1990s. For ex-
ample, although no official statistics are available, there was about 5 million tonnes
of silage dry matter produced in Brazil in 1997 (Muhlbach, 1998), and most ofthis
was whole-plant corn silage in the rapidly expanding dairy producing states.
Muhlbach also stressed the importance of machinery "rings", which are organized
by small-scale livestock producers, and harvest contractors (e.g., professional
silage-makers) to meet the forage requirements in a "year-round" silage-feeding pro-
gram in dairy production systems. Australia and New Zealand have adapted silage
technologies that are similar to those in Brazil, but one exception has been large
round bale silage, which has been a very popular "introductory" silage system for
many dairy farms with fewer than 100 cows (Kaiser & Evans, 1997). However, the
current trend in Australia and New Zealand is clearly towards precision chop
silages made in bunker, trench, or drive-over pile silos, and the increasing use of
silage-making contractors (A. Kaiser, Wagga Wagga, Australia, personal commu-
nication, 1997).

PRINCIPLES OF SILAGE PRESERVATION

The principles of silage preservation have been reviewed comprehensively


by Barnett (1954), Watson and Nash (1960), Woolford (1984), and McDonald et
al. (1991) and are discussed in greater detail in Chapters 2 and 3 of this monograph.
The major transactions that occur in silage fermentation are described briefly here
with reference to the consequent changes in nutritional value of the ensiled prod-
uct compared with the original fresh crop.
The most important compositional change is the conversion of fermentable
(water-soluble) carbohydrates-principally the hexoses (six-carbon sugars) and
fructans (fructose polymers)-to organic acids (mainly lactic acid). The basic bio-
chemical pathways are described in McDonald et ai. (1991). Since their first defi-
nition (OrIa-Jensen, 1919), lactic acid-producing bacteria are comprised of two main
groups: those that ferment hexose sugars to lactic acid alone (homofermentative)
and those that ferment hexose to lactic acid and other end products (heterofermen-
tative). The homolactic organisms reduce one molecule of either fructose or glu-
cose to two molecules of pyruvate by the Embden-Meyerhof-Parnas glycolytic
pathway and the two molecules of pyruvate then are reduced to two molecules of
lactic acid. The sum of the reaction is shown in Table 1-6 and is known as the ho-
mofermentative fermentation of sugars to lactic acid. There is no loss of dry mat-
ter as carbon dioxide and no loss of energy.
The heterofermentative organisms ferment glucose and fructose to one mol-
ecule of lactic acid and either ethanol or acetic acid and mannitol depending on the
oxidation-reduction potential of the system. Some lactic acid bacteria also can re-
duce lactate to acetate in presence and absence of oxygen (McDonald et aI., 1991).
Because only one molecule of the strongest acid, lactic acid, is produced by het-
erolactic fermentations, these bacteria are less efficient at reducing the pH of the
HISTORY 17

Table 1-6. Principal biochemical changes in silage fermentation.t

Lactic acid bacteria


Homofermentative
Glucose or Fructose + 2 ADP + 2 Pi = 2Lactate + 2 ATP + 2 H20
Losses: dry matter. none; energy. 0.7%
Heterofermentative
Glucose + ADP + Pi = Lactate + Ethanol + CO 2 + ATP + H 20
Losses: dry matter, 24%; energy, 1.7%
3 Fructose + 2 ADP + 2 Pi = Lactate + Acetate + 2 Mannitol + CO 2 + 2 ATP + H 20
losses: dry matter, 4.8%; energy, I.Oo/c
Enterobacteria
Glucose + 3 ADP + Pi = Acetate + Ethanol + 2 CO 2 + 2H2 + 3 ATP + 2 H 20
Losses: dry matter, 4.8%; energy, 17%
Yeasts
Glucose + 2 ADP + 2 Pi = 2 Ethanol + 2C0 2 + 2 ATP + 2 H 20
Losses: dry matter, 48%; energy, 0.2%
Clostridia
2 Lactate + ADP + Pi = Butyrate + 2C0 2 + 2 H2 + ATP + H 20
Losses: dry matter, 51 %; energy, 18.4%

t Sources: McDonald et aI. , 1973; McDonald et ai., 1991.

ensiled material than the homolactic bacteria. Not surprisingly, almost all silage in-
oculants contain homolactic bacteria, and the objective in adding an inoculum to
the crop is to dominate the natural mixed populations of bacteria on the crop (Lin
et aI., 1992), so that virtually the sole product of the fermentation is lactic acid.
Other groups of bacteria can develop in silage with losses of both dry mat-
ter and energy (Table 1-6). Some coliforms, or enterobacteria, can ferment glu-
cose to lactate, acetate, ethanol, succinate, and formate. Yeasts can convert glucose
to carbon dioxide and water in the presence of oxygen and can ferment glucose to
ethanol in the absence of oxygen, with substantial loss of dry matter but little loss
of energy (Table 1-6). Clostridia can ferment lactate to butyrate, with significant
loss of both dry matter and energy from the system (Table 1-6). This process is
known as secondary fermentation and also can be associated with substantial de-
carboxylation and deamination of amino acids from plant protein (McDonald et aI.,
1991). Some yeasts can utilize lactic acid and ethanol in anaerobic conditions.
With the exception of the homo lactic fermentation of glucose, the loss of dry
matter exceeds that of energy. As a result the gross energy content of silage is usu-
ally somewhat higher than that of the fresh crop, especially in wetter crops where
the extent of fermentation is greater than in drier crops (see below).
The extent and pattern of fermentation in the silo are influenced by many fac-
tors, but the main effects are those brought about by the crop itself, principally its
dry matter content, buffering capacity, water-soluble carbohydrate (WSC) con-
centration, and nitrate content (Weissbach et aI., 1974; Weissbach, 1996). The ef-
fect of increased dry matter concentration on the fermentation of Italian ryegrass
is shown in Table 1-7. In this experiment, the grass was either harvested without
wilting or wilted in the field for I or 2 d in good weather. The main effect of in-
creasing the dry matter content of the crop at harvest was a marked restriction in
the extent of fermentation, as shown by a much higher pH of the wilted silages com-
18 WILKINSON ET AL.

Table 1-7. Effects of increasing the dry matter content of Italian ryegrass by field wilting on fermenta-
tion products.t

Item Unwilted Wilted

Id 2d
Dry matter, g kg- 1 159 336 469
pH 3.7 4.1 4.9
Volatile N, gkg- 1 Total N 69 59 43
Water soluble carbohydrates, g kg- 1 DM 17 117 164
Lactic acid, g kg- 1 DM 121 54 17
Acetic acid, g kg- 1 DM 36 21 12
Butyric acid, g kg- 1 DM o o o
t Source: McDonald, 1976.

pared with the unwilted silages, a higher concentration of residual water-soluble


carbohydrates in the wilted silages, and a substantially lower concentration of lac-
tic acid in the wilted silages than in the unwilted silages.
The impact of the buffering capacity of a silage crop on the fermentation
process is illustrated best by comparison of the typical rates of acidification of en-
siled grass (relatively low buffering capacity) and legume crops (higher buffering
capacity), both of similar dry matter contents. Higher buffering capacities are as-
sociated with slower acidification and a greater risk of secondary fermentations
(Weissbach, 1996).
The buffer system in crops (weak acids and salts in equilibrium) normally is
studied between pH 6, at which many fresh crops enter the silo, and pH 4, that of
well-preserved silage. The typical ranges of buffering capacity of three contrast-
ing types of crops are shown in Table 1-8 for both the fresh crops and their silages.
The main reason for the higher buffering capacity of legumes is their higher con-
centration of organic acids (McDonald et aI., 1991).
A general association exists between the buffering capacity of the fresh crop
and that of the silage made from it. The very large increase in buffering capacity
of silage compared with that of the fresh crop is important, because neutralization
of silage acidity, once the material has been consumed by livestock, requires con-
siderable quantities of alkali, principally salivary bicarbonate.
Water-soluble carbohydrate concentration can influence the pattern and ex-
tent of the silage fermentation because acidification only can proceed if enough fer-
mentable substrate is available to the bacteria. Unfortunately, crops like legumes
typically have relatively low concentrations ofWSC as well as having relatively high

Table 1-8. Typical range of buffering capacities of different fresh crop species and their silages. t

Buffering capacity
Item Fresh crop Silage
- - - - mE kg-I of DM - - - -
Corn 200-250 900-1200
Ryegrass 250-350 1200-1500
Alfalfa (lucrne) 500-550 1750-2500
t Source: McDonald et aI., 1991.
HISTORY 19

buffering capacities, so the addition of sugar suchas molasses or removal of water


by wilting can improve the quality of their fermentation considerably (Carpintero
et aI., 1969). A "critical" concentration ofWSC exists in fresh grasses, below which
the risk of inadequate acidification is increased. In an analysis of 231 fresh crops
and the silages made from them, Wilkinson et al. (1981) found that the critical value
for WSC was 20 g kg-I fresh crop weight. Above this level, the incidence of sec-
ondary fermentations was very low.
Weissbach et al. (1974) found that the ratio between WSC (g kg-I DM) and
buffering capacity (grams lactic acid per kilogram DM required to reduce the pH
from that of the crop at harvest to pH 4.0) was a suitable measure of the acidifica-
tion potential of a crop. However, dry matter content per se can exert a marked ef-
fect on extent of fermentation (Table 1-7). Reduction of the water content of the
crop reduces water activity (the water in which bacteria can grow freely) and in-
creases the pH value at which the fermentation stabilizes (Weissbach, 1968).
Integrating dry matter concentration, WSC, and buffering capacity produces
a minimum dry matter content for stable silage in relation to crop fermentability,
or ensileability, characteristics.
The nitrate concentration of a crop can have both positive and negative ef-
fects on fermentation quality (Weissbach et aI., 1993). The positive effect is that
nitrate is reduced to nitrite in the early stages of the fermentation, and nitrite has a
strong inhibitory effect on Clostridia (Wieringa, 1966). In well-preserved silages,
a relatively small proportion of crop nitrate is reduced to ammonia, but in poorly
preserved silages, virtually all the nitrate is reduced to ammonia, which has a neg-
ative effect on fermentation quality, because the ammonia increases the buffering
capacity of the crop.
The silage fermentation affects the nutritional value of silage, as shown in
Table 1-1. Many studies have shown the intake of silage to be lower than that of
the fresh or the dried feeds made from the same original crop. For example, Waldo
et al. (1966) found that voluntary intake of silage was 0.28 kg lower than that of
hay when both were offered ad libitum to heifers. The extent of the decrease in in-
take of silage compared with hay was greater the longer the period of feeding and
greater for 7-mo-old lambs than for mature sheep (Harris et aI., 1966). Demarquilly
(1973) reported that the intakes of 87 silages by sheep were on average 33% lower
than those of the fresh crops from which they were made. However, the reductions
in intake were variable, ranging from 10 to 64%.
Much research has investigated the fermentation characteristics that might be
responsible for the lower intake of silage compared with fresh or dried materials.
Degradation of protein and amino acids to amines or ammonia, excessive acid pro-
duction, and high levels of acetic acid have been blamed for depressions in volun-
tary intake (Wilkinson et aI., 1976). These and other possible factors are discussed
in detail in Chapter 10.

GEOGRAPHIC AND CLIMATIC FACTORS THAT


AFFECT SILAGE PRODUCTION
The development of silage has been dominated by two main features: (i) an
unsuitable climate for the rapid field-drying of crops and (ii) the availability of ma-
20 WILKINSON ET AL.

terials and equipment to ensure that the crop can be harvested and contained for sev-
eral months in anaerobic conditions. In many areas of the world, for example in Sub-
Sahara Africa, the climate for much of the year is suitable for the rapid production
of field-dried hay, and equipment for silage, such as forage harvesters, is not read-
ily available. As a result, silage making is not popular in many tropical and sub-
tropical regions and in areas where the terrain is too rough for mechanized equip-
ment to be used in fields. Further disadvantages of silage in warmer regions of the
world are that the development of undesirable microorganisms is likely to be more
rapid than in cooler climates. This problem is exacerbated by the crop species en-
siled in tropical regions, and the need to achieve good consolidation of the crop in
the silo.
Wet silage poses special problems of poor fermentation quality, especially
clostridial activity and risks to cheese making from contamination of milk by
clostridial spores. In areas like northern Europe high rainfall makes field wilting
unpredictable. The AIV process (Virtanen, 1933) of adding hydrochloric and sul-
furic acids to unwilted crops at harvest to achieve immediate acidification was so
successful that today virtually all the silage made in Finland is made with an addi-
tive (Wilkinson et aI., 1996).
In other regions, the risk of secondary fermentations in silages is considered
too great and hay has remained the principal method of forage conservation. Silage
making is prohibited by law in some areas of Switzerland and Italy (Wilkinson et
aI., 1996). In these areas, high-value hard cheeses are produced, and the secondary
"blowing" of the cheese, during maturation, caused by the fermentation of the lac-
tic acid in the cheese by clostridial spores from poorly preserved silage contami-
nating the milk, can lead to the total loss of the cheese and substantial loss of in-
come to the producer as a result.
One of the major limitations of livestock production in the dry-humid and
humid tropics in the twentieth century has been the uneven supply of pasture for-
age during the year (Preston, 1982). Most of the tropics have differentiated wet and
dry seasons, and there is commonly a surplus of forage during the wet season and
a shortage during the dry season. Tropical grasses and legumes typically having
higher concentrations of cell-wall components and lower levels of WSC than tem-
perate crop species (Catchpoole & Henzell, 1971; Wilson & Ford, 1973; Jarrige et
aI., 1982), especially when harvested at immature stages of growth (Wilkinson,
1983a). In the past two decades, silage technology has been successfully introduced
into several countries in Central and Latin America and South and East Asia
(Wilkinson, 1983a; BoIsen & Faylon, 1994; Tripathi et aI., 1995). It is essential to
chop the cut forage to achieve a successful fermentation. Losses of dry matter in
small silos in the tropics can be extremely high because of poor consolidation
(Pizarro & Vera, 1980), which is partly due to the structural rigidity of tropical for-
ages and their resistance to compaction (Miller et aI., 1966) and partly due to the
lack of suitable harvesting and chopping equipment on smaller farms (Wilkinson,
1983b; BoIsen & Faylon, 1994).
A "small bag" silage system was developed recently in Thailand (Nakama-
nee, 1997). Forage is harvested by hand, chopped, and placed into a polyethylene
bag that contains 10 to 30 kg of fresh material, which is then compressed manu-
ally and sealed. Each bag provides silage for only a few "livestock days", which
HISTORY 21

reduces the risks of aerobic deterioration when the silos are opened in hot, humid
climatic conditions. Innovative approaches of this kind will be essential to get silage
into smallholder farming and livestock systems.
BoIsen and Faylon (1994) drew attention to a wide range of feedstuffs and
agricultural byproducts that may be used to improve the ensileability and/or nutri-
tive value of tropical forages. These include molasses, cereal brans, citrus pulp, and
other fruit and crop processing residues. There are also possibilities for making silage
from byproducts alone such as sugarcane bagasse and oil palm fronds (P. Zainal,
University Putra Malaysia, Serdang, Malaysia, personal communication, 1998), but
these silages are likely to have a very low nutritive value and will be useful only as
maintenance feeds.

ECONOMIC FACTORS THAT AFFECT SILAGE PRODUCTION

The increased adoption of silage by farmers in the latter half of the twenti-
eth century has reflected an appreciation that investment in silage-making equip-
ment and technology is likely to result in an acceptable return on the financial in-
vestment. For example, Rotz and Harrigan (1997) compared the production costs
of alfalfa either grazed, made into silage, or made into hay with cost of com silage
on a 100-cow dairy farm in the USA. Cost per tonne of dry matter was highest for
the hay, because machinery was used for a smaller amount of crop than for alfalfa
silage. Com silage costs were somewhat lower than those for alfalfa silage. Thus,
silage making, at least on a I OO-cow dairy farm, was financially attractive compared
with hay making. Significantly, grazing was less costly per tonne of dry matter, but
the grazing of older replacement heifers and mature cows on alfalfa with supple-
mentary feeds for up to 6 mo of the year was not associated with an increased net
return, because the same machinery and crop storage costs were depreciated over
a smaller amount of preserved forage, thus increasing the cost per tonne of preserved
forage.
The Rotz and Harrigan (1997) model and earlier simulation models (e.g., Cor-
raIl et aI., 1982) allow the effect on margins of management decisions (e.g., choice
of crop, fertilizer input, machine use, climate, cutting strategy, and livestock per-
formance) to be assessed. Such models also allow "What if?" questions to be posed
and answered.
Modeling also can allow specific questions to be answered with regard to the
economic impact of biological and technological processes involved in silage mak-
ing. For example, Rotz (personal communication, 1997) commented that the return
on investment in sealing bunker silos was between 4: 1 and 8: 1, depending on the
effectiveness of the seal and the amount of silage loss prevented (Huck et aI.,
1997). Pitt and Liebensperger (1987) developed a simulation model of the ensiling
process to study the potential benefits of inoculation of crops prior to being ensiled.
They concluded that to produce a consistent benefit in silage quality, an inoculant
should contain acid-tolerant bacteria added at levels of at least 105 g-I of fresh crop.
One area lacking in development is smaller-scale ensiling technology. The
capital investment in silage-making equipment is so large that suboptimal use of
machinery and storage can increase the cost of complementary cropping activities
22 WILKINSON ET AL.

on the same farm (Rotz & Harrigan, 1997). Economies of scale must be achieved
as far as possible to maximize the return on investment.
The potential biological benefits of ensiling, including higher digestibility and
energy value as the result of more frequent or earlier cutting or reduced losses be-
tween cutting and feeding, also can be negated if appropriate equipment is not avail-
able or not used when available. Large round or rectangular bales can be cost-ef-
fective for smaller farms, but only if wrapped and stored correctly. In developing
countries, novel techniques are required so that small holders without access to trac-
tors, forage harvesters, or balers can make silage with low storage losses and with
a nutritional value that is superior to the mature hay or crop residues (Wilkinson,
1983b). This aspect of silage development constitutes a challenge to research work-
ers that has yet to be met successfully.

ENVIRONMENTAL FACTORS THAT AFFECT


SILAGE PRODUCTION

Silage has had a poor reputation for more than 100 yr for its adverse impact
on the environment. The first report of silage effluent polluting water and making
it unfit for human consumption in the UK was made by Jenkins in 1884. Since then
the pollution of air by foul-smelling silage has made the persons responsible for pro-
ducing the silage into unpopular neighbors, who, along with farmers who allow ef-
fluent to escape into rivers and streams, have been liable to prosecution in the UK
in recent decades.
The problem of silage effluent was discussed by Woolford (1978b), who high-
lighted the relatively high biochemical oxygen demand of the material compared
with livestock wastes. The factors affecting the production of effluent and its man-
agement were reviewed in detail by Stark and Wilkinson (1988) and Offer et al.
(1991). The removal of water from the crop prior to ensiling, either through the nat-
ural process of crop maturation, as in corn, sorghum, and whole-crop cereals, or
by field wilting, is the most practical methods of reducing effluent loss from the
silos. Alternative approaches include preharvest desiccation with formic acid,
which was considered briefly in the 1970s as a possible technique for reducing ef-
fluent, but results were disappointing (McIlmoyle, 1977). More recent studies have
shown limited success with absorbent additives (Offer et aI. , 1991).
In some countries of Western Europe, the production of wet grass silages by
harvesting crops with little or no field wilting was stimulated by the Eurowilt se-
ries of experiments (Zimmer & Wilkins, 1984). This was reinforced by subsequent
work in Northern Ireland (Small & Gordon, 1988) and sustained particularly in cli-
mates that were characterized by unpredictable weather with frequent periods of
rainfall during the growing season. The losses of dry matter during wilting, in the
silo, or as effluent in the Eurowilt series of experiments were summarized by
Wilkinson (1987).
Essentially, the Eurowilt experiments showed that unwilted and wilted silages,
provided they were well-preserved, resulted in little difference in livestock per-
formance. However, Small and Gordon (1988) showed that output of milk per
hectare ofland was lower with cows fed wilted than with those fed unwilted silage.
HISTORY 23

Their work had a profound influence on silage-making practice in the higher-rain-


fall regions of Western Europe for many years. Not until the introduction of tighter
environmental controls in the 1990s did farmers start to look more closely at new
approaches to field wilting that did not involve leaving the crop exposed to the risk
of rain damage for several days .
Plant and swath limits to the rate of water loss from mown swaths were elu-
cidated in the 1970s by Jones and Harris (1980), but the full potential of their work
was realized only recently. This was the case despite the fact that the development
of a high dry matter silage system based on tedding young swaths repeatedly dur-
ing the period of field wilting was developed in Holland between 1963 and 1973
as a way of reducing the incidence of secondary fermentations in silages without
the use of acid additives (Thomas, 1977). The mower-conditioner was seen as an
alternative to the tedder, especially for high-yielding, relatively mature, grass crops.
Mower-conditioners were designed to produce relatively narrow, erect swaths to
maximize the exposure of the mown grass to evaporation of water by air movement
over and through the swath.
Farm-scale evaluation of mower-conditioners indicated that wider swaths
dried faster than narrower ones and had a greater influence on drying rate than con-
ditioning, confirming earlier work indicating that conditioning was likely to be ef-
fective only if enough energy could enter the crop to produce evaporation of water
(Dernedde, 1980). Wilkinson and Leaver (1994) and Wilkinson et al. (1996) found
that spreading the swath once immediately after mowing so that it covered the en-
tire ground area was reflected in more rapid rate of water loss and higher feed in-
take than leaving the crop in swaths during the wilting period. The finding that swath
density (weight per unit area) had a major impact on rate of water loss agreed with
research by Wright et al. (1997), who showed clearly that spreading the crop after
mowing had a greater influence on rate of drying than either conditioning or ted-
ding, and that the benefits of conditioning or tedding were increased at lower swath
densities compared with higher densities. The Dutch grass preservation system in-
volved the mowing of young crops of relatively low yield (Thomas, 1977), which,
after initial spreading over the entire ground area, would have had low densities and
rapid rates of water loss.
The recent introduction of mower-conditioners that spread the crop over the
complete area of a field should give added impetus to the rapid wilting of grasses
and legumes and thereby reduce the risk of any adverse environmental impact from
the resultant silage.

CONCLUSIONS

Silage will continue to have a substantial role in crop and livestock produc-
tion systems in the temperate areas of the world to supply energy, protein, and fiber,
especially to housed animals during periods of little or no forage growth. There will
be a need for new technologies and inputs to constrain the costs of silage produc-
tion in Europe and North America, where silage making is well established. It is
probable that there will be some replacement of perennial grass silages with silages
made from com, whole-crop cereals, and forage legumes. In contrast, in some tem-
24 WILKINSON ET AL.

perate and tropical areas, where silage currently supplies only a small proportion
of nutrients (e.g., Australia, New Zealand, and Latin America), there will proba-
bly be increases in silage production to capitalize on the advantages of a silage sys-
tem in producing a more even seasonal supply of nutrients from grassland and for-
age compared with grazing or hay systems.
The potential of silage to contribute to tropical and subtropical livestock pro-
duction is, as yet, largely unrealized. Appropriate technologies for making silage
from forages and byproducts are emerging, but there will be a continued require-
ment to harvest material of reasonable nutritional value and to maintain that feed-
ing value and reduce losses during the storage and feeding periods. There are op-
portunities for the cooperative use of chopping machines and for storage in small,
well-sealed silos. The goal of every silage maker in the past, present, and future per-
haps was best stated by E. Zimmer, at the Forage Conservation towards 2000 Con-
ference in Braunschweig, Germany, in January 1991: "to produce a silage which
has the smell of sweet perfume" .

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