Animal Feed Science and Technology, 45 (1993) 35-56 35

0377-8401/93/$06.00 © 1993 - Elsevier Science Publishers B.V. All rights reserved

Silage additives
Nancy Henderson1
Scottish Agricultural College, WestMains Road, Edinburgh EH9 3JG, UK

Abstract

The biochemical and microbiological factors involved in the ensilage process are reviewed, and the
effects of wilting and of the various categories of silage additives, stimulants, inhibitors, nutrients and
absorbents on these factors are discussed. Some examples are given of the effects of additives on dry
matter loss, especially through effluent production, and on voluntary intake and animal production.
It is concluded that given appropriate conditions relating to weather, substrate availability and good
management, a well-preserved silage may be prepared with relative ease. However, when conditions
are less favourable, the use of additives will aid fermentation.

Introduction

It is estimated that in Western Europe 60% of the forage conserved for win-
ter feed, equivalent to approximately 77 Mt of dry matter (DM), is in the
form of silage (Wilkinson and Stark, 1987). In western parts of the UK the
proportion is even higher. In Northern Ireland, for example, grass silage ac-
counts for approximately 85% of conserved forage (Mayne and Steen, 1990 ).
The conservation of a crop as silage depends upon the natural fermentation
under anaerobic conditions of the sugars in the crop to acids, mainly lactic
and acetic, by the lactic acid bacteria. Although silage is likely to represent
50-60% of the winter feed for ruminants, the silage fermentation process is
largely left to chance. Most of the soluble carbohydrates present in the fresh
forage are fermented to reduced products, which themselves may be end-
products of ruminal fermentation, and most of the nitrogenous compounds
are rendered highly degradable. Silage energy and nitrogen sources are there-
fore not synchronized for optimal utilization of ammonia by rumen organisms.
Silage additives have been developed over the years to take some of the risk
out of the ensilage process and to improve the nutritive value of silages. Ide-
ally, a silage additive should be safe to handle, reduce DM losses, improve the
hygienic quality of the silage, limit secondary fermentation and improve
aerobic stability, increase the nutritive value by increasing the efficiency of
utilization of the silage and give the farmer a return greater than the cost of
the additive (Merensalmi and Virkki, 1991 ).

~Present address: c/o Dr. F. D'Mello, Scottish Agricultural College, West Mains Road, Edin-
burgh, EH9 3JG, UK.
36 N. Henderson / Animal Feed Science and Technology 45 (1993) 35-56

Silage additives may be chemical or biological, and can be categorized as

stimulants, inhibitors, nutrients or absorbents (McDonald et al., 1991 ). The
list of commercial products available to farmers in the UK is long and di-
verse. Approximately 140 named products were on the market in 1991 (Wed-
dell et al., 1991 ), but to understand how their application to crops affects the
ensilage process it is necessary to have some knowledge of the reactions in-
volved in the process.

The ensilage process

Biochemistry

Biochemical processes occurring after the plant is cut and during conser-
vation result from the continuing metabolism of plant cells, from the enzymes
of the dead tissue and from the micro-organisms present on the plant (Table
1 ) (Henderson, 1991 ).

Respiration
Plants obtain energy and reducing power not only from light reactions of
photosynthesis but also from the degradation or respiration of products of
photosynthetic carbon dioxide fixation (Duffus and Duffus, 1984). The
overall reaction for respiration is generally represented as the complete oxi-
dation of a molecule of glucose:
C 6 H 1 2 0 6 "+'602 --~ 6 C O 2 + 6H2 O + energy
In the cut plant virtually all this energy is converted into heat. In the field the
heat is dissipated into the atmosphere but in the silo much of the heat gener-
ated is retained. Because the acceleration of respiration with temperature is
exponential there is a progressive increase with time until the oxygen supply

Table 1
Factors influencing the conservation of crops

Enzymes Micro-organisms

Respiratory Lactic acid bacteria

Proteolytic Enterobacteria
Polysaccharide-degrading Clostridia
Fungi
Yeasts
Moulds
Bacillus
Listeria
Acetic acid bacteria
Propionic acid bacteria
N. Henderson/Animal Feed Science and Technology 45 (1993) 35-56 37

is exhausted or the enzymes are inactivated by the acid conditions, or, in ex-
treme conditions, by temperatures of approximately 70 oC.

Proteolysis
In the fresh herbage 75-90% of the total nitrogen is present as protein, but
this may fall to 60-65% after ammonia or nitrate fertilization. The important
factors which influence the extent of the degradation of proteins by plant en-
zymes are DM content, the presence of oxygen, pH and temperature. During
a moist wilt with no change in the DM content there is an increase in the
amide fraction, and prolonged wilting in moist conditions increases the level
of low molecular weight nitrogen compounds, including ammonia (Spoelstra
and Hindle, 1989).
When oxygen is used up in the silo the rate of proteolysis is much more
rapid and the degradation is extensive. Plant proteases are most active be-
tween pH 6 and 7 but the activity does continue at a much reduced rate at
values below pH 4 (Heron et al., 1989). The more rapid the drop in pH in
the silo, the less extensive is the breakdown of protein. Proteolysis and amino
acid degradation are also less extensive as the DM content of herbage in-
creases, and they are inhibited by dry or wet heat treatment (Mandel et al.,
1989; Charmley and Veira, 1990).

Polysaccharide-degrading enzymes
Polysaccharides are condensation polymers based on monosaccharides
joined together by glycosidic linkages (Duffus and Duffus, 1984). Cellulose
is the most abundant naturally occurring organic compound and is the main
structural component of the plant cell wall. When grass is ensiled, lignin re-
mains unchanged and only a small decrease, less than 5%, occurs in the cel-
lulose fraction (Morrison, 1979).
Hemicellulosesare defined as a class of polysaccharides associated with cel-
lulose and are soluble in alkali. Growing primary walls in grasses are com-
posed of approximately 65% water, with 30% of the dry weight of the unlig-
nified cell wall as fl-( 1-~3), (1-~4)-glucan and 30% as arabinoxylan (Fry,
1988 ). During ensiling, losses of hemicellulose are not uniform (Morrison,
1979) and, depending on the stage of growth and DM of the grass, as much
as 40% of the hemicellulose fraction may disappear (Gonzalez-Yanez, 1990 ).
Campbell et al. (1990) concluded that pectin in lucerne is not metabolized
during ensilage but hemiceUulosecan be to a variable extent.
During the ensilage of low-DM crops the amount of acid produced is fre-
quently in excess of the water-soluble carbohydrates (WSC) in the crop. Pro-
teins, amino acids and organic acids all contribute to the production of fer-
mentation acids but the hemicelluloses are the major source of additional
substrate. Ohyama and Masaki (1977) and others have demonstrated that
much of the additional sugar produced in low-DM silages is glucose with some
38 N. Henderson /Animal Feed Science and Technology 45 (1993) 35-56

arabinose and xylose. Previously, the glucose was thought to come from cel-
lulose but now it is thought more likely that it comes from the glucan in the
primary cell walls (Fry, 1988 ). In higher-DM silages the activity of the poly-
saccharide-degrading enzymes is inhibited.

Microbiology

Lactic acid bacteria

The role of microflora in forage conservation was discussed in detail by
Woolford (1984 ) and updated recently by Pahlow (1991 ). The majority of
the bacteria on a crop require oxygen to survive and die off rapidly in the
early stages of ensilage. Of the microflora which remain, the lactic acid bac-
teria (LAB), enterobacteria, yeasts and clostridia are the most important. In
the past, it was thought that the numbers of LAB on the standing crop could
be very low. Recent research, however, has established that they are present,
but in a dormant state, and that the harvesting procedures which lacerate the
crop and release cell contents from the ruptured plant tissue result in the re-
covery of the LAB, which can be enumerated by standard methods (Pahlow,
1991 ). In general, the numbers of the epiphytic LAB on grass increase during
the summer months and may be as high as 107 colony forming units (cfu)
g-~ grass. These include the efficient homofermentative LAB which convert
glucose and fructose to lactic acid only and the heterofermentative LAB which
convert sugars to a range of products, not all of which assist in lowering the
pH. When there is no more available carbohydrate in the silage, LAB can use
lactic acid as substrate, with the production of acetic acid (Lindgren et al.,
1990). Although O'Kiely et al. (1986) concluded from a series of experi-
ments that there is a critical level of 30 g WSC 1- ~of grass juice extract below
which silages are generally poorly preserved, predicting silage pH from the
sugar content of the ensiled herbage is not possible (Mo and Fyrileiv, 1979 ).
Analyses have shown that WSC at the time of ensiling may account for only
63.3 _+25 % of the lactic acid in silage (Wilson, 1986 ).

Enterobacteria
Enterobacteria compete with LAB for available carbohydrates during the
initial stages of ensilage and some can produce ammonia (Seale, 1986 ). The
metabolic activity of the enterobacteria is readily inhibited during the conser-
vation process either by anaerobiosis or by acidification. The toxic substances
contained in the many Gram-negative bacteria are stable, however, and will
remain largely unaffected over extended periods (Lindgren, 1991 ). The final
concentration of these endotoxins will be closely related to the maximum
population reached by the enterobacteria. As yet, there is no proof, but it is
possible that the endotoxins may have a detrimental effect on the palatability
and therefore on the nutritive value of the silage.
N. Henderson/AnimalFeedScience and Technology45 (1993) 35-56 39

Yeasts
All yeasts grow well in the presence of oxygen, and their significant role in
the aerobic deterioration of silage is well established (Woolford, 1990). In
silage a subgroup develops which can compete successfully for fermentable
carbohydrate anaerobically. These are superseded by a subgroup able to uti-
lize organic acids. These organisms usually represent up to 100% of the yeast
flora in farm silages after extended storage. Silages with high yeast counts, in
excess of 105 cfu g-1, are likely to be unstable on exposure to air (Woolford,
1990).

Clostridia
Both saccharolytic clostridia and proteolytic clostridia are present in silages
as contaminants derived from soil particles. Clostridia are particularly sensi-
tive to water availability, and in very wet crops even the achievement ofa pH
value as low as 4.0 may not inhibit their growth. Clostridia play a major role
in the anaerobic spoilage of silage, and saccharolytic clostridial spores cause
problems in hard cheese production (Pahlow, 1991 ).

Moulds
Most moulds are dependent on oxygen for their growth and propagation
but even minute quantities of oxygen are sufficient to maintain the metabo-
lism of certain members of the group (Pahlow, 1991 ). Mycotoxins are found
not only in spoiled silage but also some distance from areas of visible mould-
ing (Oldenburg, 1991 ).

Silage additives

Wilting

Before the various categories of silage additives are reviewed, mention

should be made of wilting, regarded by many as an alternative to additive use.
In conditions in which a rapid wilt to 250-300 g DM kg-1 is possible, this
will be beneficial as it will reduce effluent production without having a signif-
icant effect on the nutritive value of the silage. In a collaboration programme
carried out in European research institutes, field losses and in-silo DM losses
averaged 18.6% and 17.1% for unwilted additive-treated silages and wilted
silages, respectively. Additives had little effect on losses when used on wilted
herbage but they did improve the nutritive value of the silage (Zimmer and
Wilkins, 1984).
Under good weather conditions the DM increases and the sugars are con-
centrated in the DM, but under poor weather conditions the DM content may
increase very little, if at all, and if the wilting period is extended over several
days soluble carbohydrates will be lost, protein-N contents may be reduced
40 N. Henderson/AniraalFeedScienceand Technology 45 (1993) 35-56

Table 2
Composition of'ineffective wilt' silages

Additive pH Ammonia-N WSC Lactic acid

(gkg -t total N) (gkg -~ DM) (gkg -~ DM)

None 3.82 153 0 150

ADD-F
(formic acid) 4.02 117 67 96
Silaform
(formic acid-formalin) 3.81 143 37 102
Sylade
(sulphuric acid-formalin) 3.97 144 14 115
Kylage
(calcium formate-sodium 3.95 152 1 158
nitrite)

and deamination of amino acids may increase. If this occurs the silage is likely
to have a high ammonia-N content even with the application of an effective
additive (N. Henderson and P. McDonald, unpublished data (Table 2 ) ). It
is generally accepted that a well-preserved silage should have an ammonia-N
content less than 80 g kg- 1total nitrogen (TN).

Carbohydrate sources

Carbohydrate-rich materials such as sugar, molasses, whey, citrus pulp and

potatoes are added to silage crops to increase the supply of substrate for the
LAB. Molasses is the carbohydrate source used most frequently, and is of
particular benefit when applied to crops low in soluble carbohydrates such as
legumes and tropical grasses, although to obtain maximum benefit it must be
used in relatively high concentrations (about 40-50 g kg -1 ). If the treated
crop has a very low DM content, a considerable proportion of the added car-
bohydrate may be lost in the effluent during the first few days of ensilage. The
effect of this and any other category of silage additive on DM loss between
the field and the feed trough will depend upon their effect on respiration,
effluent flow, inhibition or stimulation of the various micro-organisms and
their activities, and on the aerobic stability of the silage.

Acid-based additives

For years, acid-based additives were the most widely researched and used
in Europe and North America. Since the introduction of the AIV process
(Virtanen, 1933) in which mineral acids were applied to the crop to lower
the pH to 3.5 and, more recently, with the development of efficient additive
applicators there has been an increase in the use of mineral and organic acids,
their salts and acids mixed with formalin. Until recently, products such as
N. Henderson/Animal Feed Science and Technology 45 (1993) 35-56 41

ADD-F (formic acid) (BP Nutrition UK Ltd., Northwich, UK) and Sylade
(sulphuric acid-formalin) (ICI Ltd., Cleveland, UK) dominated the UK
market. By lowering the pH of the herbage, acids inhibit the activities of the
respiratory and proteolytic enzymes. Whether acid additives act as stimulants
or inhibitors of LAB depends upon the concentration of the active ingredient
or ingredients in the commercial product and upon the rate at which the prod-
uct is applied to the crop. Acid salts are less effective than the equivalent acid
and therefore they must be applied at a higher rate to obtain a similar effect.

Mineral acids
Mineral acids lower the pH of the herbage, inhibiting the activity of unde-
sirable bacteria such as enterobacteria and clostridia and stimulating the LAB
to use the available substrate and lower the pH further. In crops in which
substrate is in short supply this can be beneficial. Sulphuric acid is cheaper
than organic acids but its use as a silage additive has been called into question
(Mayne and Steen, 1990). Poor responses in animal performance obtained
with sulphuric acid may be related to detrimental effects on liver copper sta-
tus as observed by O'Kiely et al. ( 1989 ).

Organic acids
Organic acids, in particular formic acid, have an antibacterial action, as a
result of both a hydrogen ion concentration effect and a selective bactericidal
action of the undissociated acid (Woolford, 1984). When formic acid or sul-
phuric acid is added to grass to lower the pH to a similar level the composi-
tion of the silages may be very different (Table 3) (Carpintero et al., 1979).
In this example, formic acid restricts the activity of the LAB, thus conserving
WSC in the silage. This should increase the silage energy available for micro-
bial growth in the rumen.
Yeasts have been found to be particularly tolerant of formic acid, and high
counts of these organisms have been noted in silages treated with this additive
applied at the recommended rate (Henderson et al., 1972). Under anaerobic

Table 3
Composition of acid-treated silages

Additive Application Grass Silage Ammonia-N WSC Acetic acid Lactic acid
rate pH pH (gkg -~ TN) (gkg -~ DM) (g kg-~ DM) (g kg-~ DM)
(lt -1 )

Untreated - 5.85 3.87 95 12 28.8 122

Formic acid
(85%) 4 4.05 3.88 12 211 4.5 66
Sulphuric acid
(50%) 3 4.00 3.64 42 64 20.4 102
42 N. Henderson/Animal Feed Science and Technology 45 (1993) 35-56

conditions yeasts obtain energy from the fermentation of sugars with the pro-
duction of ethanol and loss of DM.
As enterobacteria are known to generate formic acid, this acid probably has
less effect on reducing their growth than the growth of LAB, and it is therefore
the rapid proliferation of LAB rather than the decrease in pH associated with
the application of formic acid which is important in reducing numbers of
enterobacteria (Chamberlain and Quig, 1987). Intermediate levels of appli-
cation of formic acid ( 3-41 t - 1) may inhibit the LAB to a greater extent than
the enterobacteria and thus have a deleterious effect on the fermentation. At
higher levels of application both LAB and enterobacteria are inhibited. Formic
acid is frequently used as a positive control treatment in experiments de-
signed to evaluate other silage additives. The rate at which the formic acid is
applied may therefore have a significant effect on the conclusions drawn from
the results of these experiments.
When Mayne and Steen (1990) investigated the data from 21 recent stud-
ies in which the effects of using formic acid on fermentation and animal per-
formance were considered, they concluded that the results supported the hy-
pothesis of Parker and Crawshaw (1982) that in situations where treatment
with formic acid resulted in an improvement in silage fermentation, positive
effects on digestibility and intake of silage were obtained, and that these ef-
fects were reflected in enhanced animal performance. However, acid addi-
tives can increase effluent production on young grass by up to a third depend-
ing on the level applied (McAllan et al., 1991 ).
When formic acid is applied at a high level ( 5 1 t - 1 or more) much of the
WSC is retained in the silage, and the acid content and buffering capacity are
much lower than those of an untreated silage from the same sward (Hender-
son et al., 1990a). Some of the data from a trial conducted at Edinburgh are
shown in Table 4 (Henderson et al., 1989). In this experiment, application
of formic acid was sufficient to lower the pH of the grass immediately to 4.0.
The beneficial effects of this are seen in the improved intakes and perform-
ances of the steers given the treated silage supplemented with 1.5 kg of brew-
ers' dark grains per day compared with those of the steers on the untreated
silage with the same supplement.

Formaldehyde-acids
Interest in formaldehyde as a silage additive arose from its bacteriostatic
properties and because it was known to protect plant proteins from degrada-
tion in the silage and in the rumen. When applied at high levels, formalin
depresses DM digestibility and intake, whereas at low levels of application it
tends to encourage growth of clostridia. For this reason, commercial products
contain mixtures of acids and formalin. The effects of formic acid and a formic
acid-formalin additive on the composition and nutritive value of ryegrass
silages are shown in Table 5 (Hinks et al., 1980). In addition to preserving
N. Henderson/AnimalFeedScienceand Technology45(1993)35-56 43

Table 4
Composition and nutritive value of silage treated with a high level of formic acid (5 1t - 1 )

Treatment Untreated Formic acid SED

DM (g kg -a) 224 233

pH 3.70 3.91
Ammonia-N (g kg- ~TN) 69 35

Components of DM (g kg- l)
WSC 18 105
Lactic acid 137 51
Digestible organic matter (DOM) 718 719
Metabolizable energy (ME) (MJ kg -~ 11.4 11,7
DM)

Intake and liveweight gain (LWG) of steers

Silage DM intake (kg day- ~) 6.00 6.85
Total DM intake (kg day-~ ) 7.33 8.18 0.145
LWG (kg day -1) 0.658 0.804 0.0431

Table 5
Composition and nutritive value of wilted silages treated with either formic acid or a formic acid-
formalin mixture

Treatment Formic acid Formic acid plus formalin SEM

(3.14 kgt - t ) (2.86 kg acid t -~
plus 1.44 kg formalin t -~ )

DM 272 280
pH 4.09 4.08
Protein-N (g kg -~ TN) 383 508
Ammonia-N (g kg- ~TN) 70 50

Components of DM (g kg- 1)
WSC 57 99
Lactic acid 92 74
ME (MJ kg -~ DM) 10.4 11.1

In takes and L WG of steers

Silage DM intake (kg head- ~) 764 765 32.4
Total DM intake (kg head -~ ) 895 892 33.0
LWG (kg day- t )
(adjusted to constant ME intake) 0.829 0.933 0.0516

WSC, additives based on acids, especially those containing formalin, inhibit

proteolysis by lowering the pH and through the binding of formaldehyde to
the nitrogenous components (McDonald et al., 1991 ). This, in turn, reduces
the deamination of the amino acids and the production of ammonia.
Although formalin-acid mixtures are very effective on grass, their use has
been banned in some countries. The potential health risks of exposure to
44 N. Henderson/AnimalFeedScience and Technology 45 (1993) 35-56

formaldehyde are now recognized, and effort should be put into the contain-
ment of formaldehyde vapour where it is used.
Acid salts
Despite the positive effects of acids, especially organic acids, their corro-
sive action against machinery and their health risk towards man if precau-
tions are not taken in handling them have focused attention on alternatives
such as acid salts.
The inhibitory effect of nitrite on clostridia in silages has been studied in
detail (Spoelstra, 1985 ), and nitrite-containingadditives are marketed in most
countries in Western Europe, although up to now results with these additives
have been variable. In an experiment in which grass was ensiled after a long
ineffective wilt, a commercial additive containing sodium nitrite and calcium
formate was less effective than acid additives when added at the recom-
mended rate (Table 2 ).
To overcome the problems associated with formic acid, a complex acid salt,
ammonium tetraformate, was developed for commercial use (Drysdale and
Berry, 1980). More recently, a product containing ammonium salts of formic
acid and propionic acid with caprylic acid, but mainly a complex salt of formic
acid, has been introduced to the UK market (Maxgrass; BP Nutrition UK).
This has a recommended rate of application of 6 1 t - 1 and will inhibit the
activity of the micro-organisms, as seen from the results of a laboratory trial
in which ryegrass was treated with this additive (Table 6) (McGinn et al.,
1990). Much of the ammonia in the treated silage was applied to the grass in
the additive and is not a product of the deamination of amino acids in the
silage.
Biological additives
Biological additives are safe to handle. They either provide additional sub-
strate for the indigenous population of micro-organisms or increase the pop-
Table 6
The composition of ryegrass silage treated with a range of commercial additives

Treatment Rate of pH WSC Ammonia-N Lactic acid

application (g kg -~ D M ) (gkg -~ TN) (gkg -~ D M )

None 0 4.05 29 90 153

Maxgrass 6.01 t -~ 4.33 184 52 (15) a 3
Molasses 9.01 t -~ 4.02 42 73 121
ADD-F 2.51 t-~ 3.92 43 29 99
ADD-F 5.01 t -~ 4.11 173 16 9
Biomax 106 LAB g-~ 3.96 34 80 141
Silaform 2.51 t-~ 3.94 93 34 92
SED 0.026 9.3 2.8 8.4

aValue corrected for ammonia in additive.

N. Henderson/Animal Feed Science and Technology 45 (1993) 35-56 45

ulation ofhomofermentative LAB. In some products, the LAB are added with
substrate or with enzymes to provide additional substrate.

Bacterial inoculants
The concept of applying strains of homofermentative LAB to herbage to
improve the fermentation is not new. Throughout this century, scientists have
experimented with the addition of bacteria with or without sugar. Whitten-
bury ( 1961 ) defined the criteria which a potential organism should satisfy
for use in silage, but it was not until freeze-drying and encapsulation tech-
niques were developed that the commercial exploitation of cultures of LAB
as additives for silage was possible. Initially, many products did not contain
sufficient live organisms to outnumber the indigenous population and domi-
nate the fermentation, but products have improved, and, provided the correct
storage conditions are maintained and the directions for use are followed,
most will give the intended inoculation rate of 105-106 cfu g- ~ herbage. Sev-
eral products contain fewer LAB but are supplied with a broth in which the
farmer grows up the LAB. If this is done at the correct temperature, around
25 °C, and with attention to detail, a study on farms in the Edinburgh area
has shown that the correct inoculation rate will be achieved.
To be successful, the LAB in the inoculant must outnumber the epiphytic
population of LAB. Pahlow ( 1991 ) found that an inoculation factor (IF) of
two, i.e. a two-fold increase in LAB, was the minimum required to achieve a
positive effect on fermentation quality, but Satter et al. (1987) ranked their
production trials according to the IF and concluded that a positive response
in milk production had only been obtained with an IF of 10 or more. When
the results of the Eurobac Conference (Lindgren and Pettersson, 1990) were
compiled, it was found that in laboratory studies successful inoculation in-
creased the lactic acid/acetic acid ratio by both increasing the lactic acid and
decreasing the acetic acid contents, lowered pH and ammonia-N concentra-
tions and decreased DM losses by 20-30 g kg-~ DM. The increased rate of
fermentation with inoculation results in the suppression of proteolysis and
deamination of herbage protein (Heron et al., 1987 ) and in a more efficient
use of WSC, with more sugar retained in the silage (Gordon, 1989 ). The data
from one trial in which a mixture of Lactobacillusplantarum and Pediococcus
pentosaceus or L. plantarum only were applied to perennial ryegrass are shown
in Table 7 (Henderson et al., 1990c). Pediococci or streptococci are included
in many commercial products, as they are active within the pH range 5.0-6.5,
but other products contain only L. plantarum, which is known to satisfy most
of the criteria suggested by Whittenbury. Although in this trial the composi-
tions of the untreated silage and silage treated with the lower rate of LAB were
similar, a time course study showed that the pH fell more rapidly in the treated
silage. Despite differences in the chemical compositions of the two silages
46 N. Henderson/AnimalFeedScienceand Technology45 (1993) 35-56

Table 7
Composition and nutritive value of silages treated with formic acid or LAB

Treatment Untreated Formic acid L. plantarum L. plantarum plus SED

(31t -~) (105cfug -1) P. pentosaceus
(106cfug - l )

DM (gkg -~) 168 182 163 181

pH 4.55 4.44 4.40 4.09
Ammonia-N (g kg- l TN ) 130 109 131 88

Components of D M (g k g - t )
WSC 0 11 0 20
Lactic acid 59 51 71 84
Acetic acid 46 35 45 30
Ethanol 13 33 9 7
DOM 710 726 723 737
ME (MJ kg -1 DM) 11.4 11.9 11.5 12.5
DM loss (%) 17.8 18.3 15.3 13.6

Intakes and L WG of lambs

Silage DM intake (g day- l ) 681 692 753 792 34.3
Total DM intake (g day-1 ) 857 868 929 968 41.8
LWG (g day- 1) 71 94 124 129 14.8

inoculated with LAB, their nutritive values were similar and significantly bet-
ter than those of the untreated or formic acid treated silages.
Since 1986, many trials have been carried out with bacterial inoculants un-
der farm conditions, and a number of these have been reviewed by Mayne
and Steen (1990). In these studies, inoculant treatment did not improve the
fermentation characteristics of the silages significantly. Despite this, re-
sponses in both silage digestibility and animal performance were obtained in
a number of studies. In nine studies in which liveweight gains were recorded,
a mean increase of 17.9% was obtained compared with untreated silages. A
much smaller response of 3.7% was obtained in milk energy output (mean of
seven studies), equivalent to an increase in milk energy output of 2.5 MJ
day- 1. Both Spoelstra ( 1991 ) and Mayne and Steen (1990) have concluded
that the content of WSC in the herbage has limited use in predicting the re-
sponse to inoculation. Even in situations where there is little improvement in
fermentation characteristics measured, the use of a bacterial inoculant can
result in an improvement in animal performance (Thomas et al., 1991 ). The
data in Table 7 (Henderson et al., 1990c) support the conclusion of Spoelstra
( 1991 ) that it is the first stage of silage fermentation and the effects of addi-
tives during this stage which require further investigation to explain how bac-
terial inoculants are influencing intake and animal performance, but some
way of measuring this in the silage must be found.
N. Henderson/AnimalFeedScience and Technology45 (1993) 35-56 47

Cell wall degrading enzymes

The use of cellulolytic and hemicellulolytic enzymes as silage additives has
been considered from two points of view; first, as a means of increasing the
content of WSC as substrate for the LAB, and, second, as a method of im-
proving the digestibility of the organic matter of the crop (McDonald et al.,
1991 ). Most commercial enzymes are crude preparations containing many
enzyme activities. As much of the hemiceUulose (up to 40%) is degraded
during silage fermentation (Gonzalez-Yanez, 1990), it is the action on cellu-
lose, or on the acid detergent fibre (ADF) fraction, which produces most of
the additional substrate (Henderson et al., 1991 ).
Enzyme preparations, like plant cell wall degrading enzymes, are most ac-
tive in immature, low-DM silages and less active in wilted and mature silages
(Spoelstra, 1991 ), and they are active over a wide temperature range (20-
50 ° C). Laboratory studies with enzyme-treated silages have shown that lactic
and acetic acid concentrations are higher than those in untreated control sil-
ages, and ammonia-N concentrations and pH are lower (Rauramaa et al.,
1987). When poorly fermentable grass is ensiled, the application of enzymes
does not prevent a butyric acid fermentation. Enzymes applied at commercial
dosages do not appear to liberate sufficient additional sugar during the onset
of silage fermentation (Honig and Pahlow, 1990 ).
Although cell wall degrading enzymes do increase the WSC content and
lower the fibre content during ensilage, in none of the animal trials reviewed
by Spoelstra ( 1991 ) was there a significant improvement in digestibility with
treatment (Jacobs and McAllan, 1990). Van Vuuren et al. (1991) demon-
strated that when dairy cows "consumed enzyme-treated silage there was a
higher rate of fermentation in the rumen and an increased outflow of organic
matter from the rumen, but these had no effect on the intake of organic mat-
ter. Improved milk production was reported by Chamberlain and Robertson
(1989) when feeding a low-concentrate diet but not when feeding the same
enzyme-treated silage in a high-concentrate diet.
Many commercial inoculants contain some cell wall degrading enzymes but,
as the optimum pH of the enzymes is 4-5, it is unlikely that they produce
sugar at a sufficiently early stage to be effective or that they are there in suf-
ficient quantities to be effective at a later stage in the fermentation. The data
from a trial carried out by Gonzalez-Yanez et al. (1990) demonstrate that it
is the bacterial inoculants which are effective in such products, rather than
the enzymes (Table 8 ). In this trial, the commercial enzyme was effective in
producing additional substrate for the LAB, principally from the ADF or cel-
lulose-lignin fraction. This resulted in a significant decrease in the digestibil-
ity of the organic matter and no significant improvement in animal perform-
ance. The silage treated with the commercial inoculant and enzymes was used
most efficiently by the lambs for liveweight gain.
McAllan et al. (1991) studied the effect of various additives on effluent
48 N. Henderson/AnimalFeedScience and Technology45 (1993)35-56

Table 8
Composition and nutritive value of enzyme-treated silages offered to lambs

Treatment Untreated Cellulase- Bacterial SED

hemicellulase inoculant-
enzyme cellulase-
hemicellulase

DM 180 202 169

pH 4.20 3.72 4.00
Ammonia-N (g kg- ~TN) 87 61 83

Components of DM (g k g - l)
WSC 3 31 6
Lactic acid 70 110 99
Acetic acid 62 32 48
Neutral detergent fibre 471 425 491
Acid detergent fibre 286 246 304
Hemicellulose 185 179 187
DOM 691 656 676 8.6

Intake and L WG of lambs

Silage intake (g DM day- ~) 785 770 811 36.0
Liveweight gain (g day- l ) 72.0 81.8 95.6 14.38

production and found that enzyme additives had the greatest effect, increas-
ing effluent flow compared with untreated grass, for both young and mature
grasses. Research is being carried out using genetic engineering to create lac-
tobacilli with the ability to degrade plant cell walls (Hahts and Javorsky,
1991).

Aerobic deterioration inhibitors

As yeasts play an important role in the aerobic deterioration of grass sil-

ages, potential deterioration inhibitors must act against yeasts (Woolford,
1990). Yeasts develop during wilting and when oxygen infiltrates the silage
during the storage period. Complete exclusion of air from a farm silo is prac-
tically impossible, but aerobic deterioration of silage can be minimized if the
silo is filled rapidly, sealed efficiently and good management is practised at
feed-out. Some additives will improve aerobic stability by delaying the onset
of deterioration, but they will not prevent it.

Acids
Propionic acid inhibits most but not all of the organisms responsible for
silage deterioration, but only when applied to crops in relatively high concen-
trations. Similarly high levels of formic acid may delay the onset of deterio-
ration. In a recent trial (Henderson et al., 1990b), perennial ryegrass was
N. Henderson/AnimalFeedScience and Technology45(1993) 35-56 49

treated with Maxgrass at a rate of 6 1 t- ~ and ensiled in bunker silos of 100 t

capacity. The chemical and microbiological composition of core samples taken
from the untreated and treated silages before and after exposure to air in poly-
styrene containers for 4 days is shown in Table 9. Lactate assimilating yeasts
increased in numbers in both silages but those in the untreated silage had a
higher capacity for lactic acid assimilation and, in the short term, the treated
silage was more stable than the untreated silage.

Bacterial inoculants
When Pahlow (1982) ensiled prewilted grass with an inoculum of LAB the
inoculant restricted the development of yeasts and rendered the silage more
stable than an untreated silage. However, reductions in stability with inocu-
lation have been reported frequently (Spoelstra, 1991 ). Inoculation can only
improve aerobic stability if yeasts are kept below the threshold value of l0 s
g-~ silage and the air supply remains low, but it is not a reliable measure
against aerobic deterioration (Honig, 1990).

Bacterial inoculants-chemicals
In an attempt to prevent inoculation with LAB producing silages with high
contents of lactic acid and very low contents of volatile fatty acids and there-
Table 9
Chemical and microbiological composition of Maxgrass-treated silage before and after 4 days' expo-
sure to air

Treatment Untreated Maxgrass (6 1 t - ~)

Before After Before After

DM (g kg - I ) 190 169 200 195

pH 3.83 8.24 3.97 4.55
Ammonia-N (g kg - 1 T N ) 89 101 111 113

Components of DM (g kg- l)
WSC 0 11 177 115
Lactic acid 132 0 51 23
Acetic acid 32 6 18 58

Micro-organisms (cfu g - 1)
LAB 7.1 X 10 8 8.0X 10 9 1.5X l0 II 3.9× 1012
Enterobacteria 2.2)< 105 1.0× 106 3.3× 104 8.9× 102
Total yeasts 5.1 X 105 1.4X 10 9 5.6× 10 s 9 . 5 × 101°
Lactate assimilating yeasts 1.1 X 107 2.1 X 101° 5.5 × 106 9.5 × 10~°
Moulds 2.5 × 102 < 10 2.3 × 10 4 4.1X 102

Temperature rise above

ambient in 4 days 15.5 ° C 0 °C
50 N. Henderson/AnimalFeed Science and Technology 45 (1993) 35-56

fore prone to aerobic deterioration, the inoculum of LAB is now being com-
bined with a chemical such as calcium formate or sodium formate (Set~il~iet
al., 1990; Weissbach et al., 1991 ). Although this approach is still at the exper-
imental stage, it appears that these salts develop an antimicrobial effect with
increasing acidity in the silage. The treated silages contain less lactic acid,
fewer clostridial spores and are more stable than corresponding untreated
silages.

Nutrients

Nutrient additives are defined as substances which, when added to ensiled

material, contribute significantly to the nutritional needs of animals consum-
ing the silage. These include molasses, cereals and whey, which also act as
fermentation stimulants (McDonald et al., 1991 ). The crude protein content
of crops such as maize which are nutritionally deficient in nitrogen can be
increased by the application of urea or ammonia. Maize is also a poor source
of calcium, and improvements in animal performance have been noted on
silages treated with limestone and urea compared with silages treated with
urea only.

Absorbents

The shift from hay to silage over the past 40 years and the realization that
direct-cut silage treated with an effective additive is used more efficiently by
stock than wilted silage (Zimmer and Wilkins, 1984) has meant that silage
effluent pollution is now a major environmental problem, and in some years
it is the predominant source of agricultural pollution (Offer et al., 1991 ).
Absorbents will alleviate the problem but in wet climates they will not elimi-
nate it, and farmers must ensure that their silos do not leak and that their
effluent tanks have sufficient capacity for the weight of grass the silos will
hold. A heavy fine or even imprisonment may result if effluent finds its way
into a nearby water course.
For environmental reasons, a rapid wilt to 250-300 g DM kg- 1 should be
the technique adopted in the future. However, heavy swaths and poor cli-
matic conditions frequently render this impossible in northern Europe. Where
there is a risk of pollution, additives, such as enzymes or formic acid, which
increase effluent flow or alter the pattern of effluent flow should be avoided,
and the use of absorbents should be considered. McAllan et al. ( 1991 ) dem-
onstrated that enzyme treatment and formic acid increased effluent flow but
bacterial inoculants had no effect compared with untreated grass. In a trial
conducted by Kennedy and Carson (1991), Maxgrass-treated silage pro-
duced 30% more effluent than untreated silage in the first 5 days.
Although some absorbents may have only a minimal effect on total effluent
N. Henderson/Animal Feed Science and Technology 45 (l 993) 35-56 51

production, they may change the pattern of effluent flow and ease the short-
term problem of effluent storage and disposal (O'Kiely, 1990a). Of the ab-
sorbents tested, fibrous by-products such as sugar beet pulp (O'Kiely, 1990b;
Ferris and Mayne, 1990) or distillers' dried grains appear most promising.
Chopped straw is an effective absorbent but it increases the silo capacity re-
quired and lowers the ME value of the combined product. Alkali-treated straw
cubes are available commercially as an alternative to chopped straw.
Under farm conditions, effluent retention rates rarely exceed 1 1kg- ~added
absorbent, and for low-DM grass the absorbent may compose as much as 38%
of the food on a DM basis for complete effluent retention. This would be
uneconomic. In the majority of animal trials, effluent production is reduced
with absorbents (Kennedy, 1988; Jones and Jones, 1988) but the effluent can
have a higher DM content than the effluent from untreated silage (Kennedy,
1988).
When Ferris and Mayne (1990) examined the effect of inclusion of sugar
beet pulp with grass at ensiling, the inclusion of increasing levels reduced ef-
fluent output from 242 1 t-~ grass ensiled in the untreated to 26 1 t-~ at the
highest level of inclusion. Although dairy cows consumed more of the ensiled
blends, there was a trend for yields of milk, milk fat and milk protein to be
greater when sugar beet pulp was offered as a supplement to control silage
rather than as an ensiled blend.

Economic appraisal of additive use

Silage additives are an insurance policy against the production of poorly

preserved, inedible silage and this is how they are regarded by many farmers,
especially dairy farmers. Additives cost from 40p to treat a tonne of herbage
with sulphuric acid up to £ 12.20 to treat a tonne of low-DM herbage with an
absorbent (Weddell et al., 1991 ). Whether or not an additive has been cost-
effective is impossible to gauge without an untreated control and detailed in-
formation on DM recovery and animal performance.
If additives do improve animal performance without significantly affecting
DM losses or DM intakes they will be cost-effective if the return exceeds the
cost of the additive. Of the fermentation inhibitors, Maxgrass is probably the
most expensive. With this type of treatment more silage is consumed by stock,
and the cost of the extra silage and additive must be included in the calcula-
tion. Savings can then be made by reducing the intake of concentrate. In a
recent trial at Edinburgh (C.A. Morgan et al., unpublished data) Maxgrass-
treated silage was compared with a control silage as a basal forage for preg-
nant ewes. Supplementary concentrate in late pregnancy was adjusted to
maintain equal ME intake and fl-hydroxybutyrate blood levels when the sil-
ages were offered ad libitum. The application of Maxgrass saved 300 g of
concentrate per ewe per day in the last 38 days of pregnancy with no signifi-
52 N. Henderson /Animal Feed Science and Technology 45 (1993) 35-56

cant differences at lambing in ewe weight, condition score or lambing per-

formance. Using costs of £ 160 t - 1 for the concentrate, £20 t - 1 for the control
silage and £24 t - 1 for the Maxgrass-treated silage, the financial implications
are shown in Table 10. The DM losses from the silages were similar (control
18.5%, Maxgrass 17.8%).
Absorbents are the most expensive of the silage additives, but inclusion of
molassed sugar beet pulp or distillers' by-products has been shown to improve
silage DM intake and animal performance when compared with untreated
silage. This strategy has the advantage that nutrients that would be lost in the
effluent are fed to stock and, in some trials (Dulphy and Demarquilly, 1976;
Jones and Jones, 1988), but not all (O'Kiely, 1990a; Steen, 1991 ), improved
performance has been observed when the absorbent is ensiled with the grass
rather than offered at the same level with untreated silage. Although the ab-
sorbent must be purchased earlier in the year, this may prove to be cost-
effective.

Conclusions

Given the right conditions, good weather, sufficient substrate for the LAB
and good management, it is possible to make a well-fermented silage. The
crop should be mown when it has reached its driest point in the day, normally
in the afternoon, and wilted for no more than 24 h. When conditions are less
than ideal, there is a wide range of effective products which will aid the fer-
mentation. In future research, the emphasis should be on the development of
additives which will reduce losses during storage, improve the efficiency of
utilization of silages, i.e. improve animal performance without increasing sil-
age DM intake, and on matching concentrates to silage type.

Table 10
Evaluation of Maxgrass-treated silage for pregnant ewes

Treatment Untreated Maxgrass

(61t -1)

Intake per ewe in 38 days (kg)

Silage 154 176
Concentrates 22.8 11.4

Cost per ewe in 38 days (p)

Silage 308 422
Concentrates 365 182
Total 673 604

Saving per ewe 69p

N. Henderson/Animal Feed Science and Technology 45 (1993) 35-56 53

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