Biomass Productivity of Mixtures

BIOMASS PRODUCTIVITY OF MIXTURES
6. R. Trenbath'
Waite Agricultural Research Institute, University of Adelaide,
Adelaide, South Australia
I.Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
11.Comparison of Yields of Mixtures and Monocultures . . . . . . . . . . . . . . . . . 179
111.Theoretical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
IV. Types of Interaction Causing Nontransgressive Deviations of Mixture
Yields from Mid-Monoculture Values .............................. 186
V. Mechanisms Capable of Causing Transgressive Yielding by Mixtures . . . . 196
VI. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
I. Introduction
Mixtures of field crops are still extensively grown in primitive agricul-

ture, but where more advanced methods are used, monocultures are usual.
Claims have frequently been made, however, that communities with some
degree of genotypic heterogeneity have advantages over pure stands. The
alleged advantages have included one or more of the following: higher
yields, lower variability of yield from season to season, a better spreading
of production over the growth period, less susceptibility to disease or lodg-
ing, and an improved quality of the crop product.
The yield advantage that would justify the growing of a mixture has
commonly been thought to be a higher yield from the mixture than from
an equal area divided between monocultures of the components in the same
proportion as that in which they occur in the mixture (e.g., Jensen, 1952).
Numerous reports of this kind of mixture advantage [reviewed in Jensen
(1952) and in Simmonds ( 1962)] gave the impression that mixed cropping
was highly desirable. More recently the advantage required to justify
mixed culture has been realized to be a superiority of the yield of the mix-
ture over that of the better (or best) of its components grown in monocul-
ture over the whole of the same area (Donald, 1963; Frey and Maldonado,
Present address: Research School of Biological Sciences, Australian National
University, Canberra City, Australia.
177
178 B. R . TRENBATH
1967). Judged from this standpoint, the performance of mixtures seems

much less encouraging. Indeed, mixtures occasionally yield less than the
less (or least) productive monoculture, a type of result which tends to
receive little emphasis.
Although a yield and quality advantage has been firmly established for
legume-grass mixtures grown for forage, the experimental substantiation
of claims for other combinations, such as varieties of field crops, has proved
difficult. This has been due to the usually small size of the advantage and
its apparent dependence on relatively specific co.mbinations of biological
material and environment. However, the possibility of gaining benefits from
the simple mixing of seeds holds such attractions for agriculturalists that
research into the productivity of mixtures continues. There are three main
objectives in this research: the first is to screen mixtures composed of
more-or-less randomly selected genotypes for “lucky,” high-yielding com-
binations. The second objective is to test the alleged advantages of tradi-
tionally grown mixtures, particularly those of some less-developed coun-
tries. The third objective is to gain an understanding of the processes which
lead to mixture advantages so that in a specific environment, a rational
choice of components may produce mixtures showing benefits unobtainable
from pure cultures.
Since there is much literature concerning the yields of mixtures (re-
viewed by Aiyer, 1949; Simmonds, 1962; Baldy, 1963; Donald, 1963),
the present survey treats only a limited part of what has been written on
this subject. The first limitation refers to sources: the reports considered
have mostly appeared in English-language and West European journals
and dissertations. The second limitation is that grazed crops are not con-
sidered although crops cut for fodder are included. A third restriction refers
to experimental design: in order to be included in the section concerned
with yields, reports must have described experiments in which biomass
(dry-matter yield of shoots) was measured in mixtures of pairs of com-
ponents planted simultaneously in 1 : 1 proportions, and in which monocul-
tures of the components were grown at the same total density within the
same experiment. The numerous reports of experiments involving mixtures
of cereal varieties (see Simmonds, 1962) mostly consider only grain yield;
since the degree of correlation between grain yield and shoot weight is
uncertain, reference will be made to such reports only where a point cannot
be adequately illustrated by data on biomass. The subject of mixtures of
legumes and nonleguminous species is not considered because a special
nutrient relationship is involved in the interaction between the components;
this type of interaction seems to be already much better understood than
many of the types of interaction occurring in the mixtures to be considered
in the present work.
BIOMASS PRODUCTIVITY OF MIXTURES 179
Apart from a section on underlying theory, this review will consider

evidence concerning the following topics : the biomass yields of mixtures
compared with those of their components’ monocultures, mechanisms caus-
ing minor (nontransgressive) deviations from the “mid-monoculture” yield
midway between the yields of the components’ monocultures, and mecha-
nisms causing transgressive deviation, so that the yield of the mixture is
either less than that of the lower-yielding monoculture or more than that
of the higher-yielding monoculture.
II. Comparison of Yields of Mixtures and Monocultures
To facilitate description, a number of terms and symbols are introduced.

The mid-monoculture yield is denoted by P. A mixture will be said to
have “overyielded” when the mixture biomass, M , has exceeded that of
the more productive pure culture, P I ; i.e., when M > P,. The mixture will
be said to have “underyielded” when the mixture biomass has fallen below
that of the less productive pure culture, P,, i.e., when M < P,. These two
“transgressive” situations will be shown to have occurred with a relatively
low frequency compared with the cases where P , > M > P,. Since an
understanding of the reasons for transgressive yielding by mixtures may
enable the agriculturalist to produce overyielding at will, attention has been
particularly focused on reports of its occurrence.
In his review of the performance of mixtures, Simmonds (1962) was
concerned almost entirely with the grain yield of cereals. His conclusions
are of interest, however, because they are so similar to those of Donald
( 1963), who considered yields of biomass. Simmonds found that often
M = P, sometimes P, > M > P, and occasionally M > P,. He noted that
“negative” interactions (i.e., M < P ) seemed to be rare.
Reviewing evidence concerning the possibility that mixtures might over-
yield in biomass, Donald (1963) concluded that the yields of mixtures
usually lay between PI and P,, and that there was no substantial evidence
to show that a mixture of two genotypes can fix more carbon than the
more productive of the two genotypes grown in monoculture. Van den
Bergh (1968) similarly concluded that in almost all experiments, the mixed
culture yielded less than the monoculture of the more productive species.
In the same way neither Donald (1963) nor van den Bergh (1968) found
strong evidence of underyielding.
To show the relationship between the observed yields of binary mixtures
and the yields of their components grown in monoculture, the results of
344 mixtures are summarized in Table I. The data are taken from pub-
lished reports of experiments in which the experimental design known as
180 B. R. TRENBATH
TABLE I
The Distribution of Biomass Yields of Varietal or Interspecific Mixtures
Compared with Yields of Their Components’ Monocultures, Based on
Published Data of 344 Mixtures”
Below Above
Crop Pz PftoP P t o p 1 PI Author
Grasses 3 Ahlgren and Aamodt

(1939)
Either grasses or leguniesc 3 1 2 Aberg et al. (1943)
1 1 9 3 Donald (1946)
Wheat 2 1 Sakai (1953)
Rice 9 12 11 4 Sakai (1955)
A series of nonlegumes~ 1 1 10 3 Williams (1962)
Subterranean clover 1 1 1 Williams (1963)
Flax and linseed 9 7 8 12 Harper (1965)
Grasses“ 12 38 42 a8 England (1965)
Barley 1 4 5 Norrington-Davies
(1967)
Grasses“ 5 9 11 15 Norrington-Davies
(1968)
Grasses 2 2 1 1 Rhodes (1968b)
Grasses“ 2 7 3 4 Thomson (1969)
Grasses 1 4 5 a Rhodes (1970a)
Rye 4 4 16 7 nor rington-Davies
and Hutto (1972)
- - - -
45 92 124 83 = 344
62.8%
c _ _ _ _T_I
39.8% 60.2%
a P I and PZ are the yields of the higher- and lower-yielding monocultures, respectively.
13 is the mid-monoculture yield.
” Data derived from a series of cuts.
c Mixtures of leguminous and nonleguminous species have been omitted.
a “mixture diallel” was used. Experiments conforming to this design con-

tain a series of genotypes grown in monoculture and all possible binary
combinations. All plots (or containers) are sown with a constant overall
density of plants, and 1 : 1 proportions are used in the mixtures. Such mix-
ture experiments satisfy the conditions specified earlier for consideration
in this part of the review. Although replication is not specifically mentioned
by Harper (1965), it is believed that all data used in Table I were based
on replicated experiments. The data of Jacquard and Caputa (1970) were
reported as being unreplicated and are therefore not included. Taken to-
gether, the data show that mixture yields tend to lie above the mean yields
of the monocultures; the difference between 60.2% and 39.8% is highly

significant ( P < 0.001 ). Presumably reflecting this difference, the reported
frequency of overyielding is significantly ( P < 0.001) greater than that
of underyielding. According to these results, apparently transgressive yield-
ing is by no means uncommon, comprising 37% of all the mixtures con-
sidered. Although differences between overyielding mixtures and their
higher-yielding monocultures have often not been examined for statistical
significance, of those so examined, very few have reached even the 5 %
probability level.
As far as the author is aware, there are only four investigations where
overyielding has been reported as being statistically significant. The first
case (Whittington and O’Brien, 1968) concerns the “significant” overyield-
ing by three mixtures of grasses in the third year of a field experiment.
This overyielding affected only the treatment with frequent cutting, but
there was also a tendency toward it in the less-frequent cutting treatment.
It should be noted, however, that the statistical significance of the over-
yielding was tested using selected data (6 out of 15 harvest periods); van
den Bergh ( 1968) has observed that apparently highly productive mixtures
in some cuts may be quite the reverse in other cuts. Since the mixtures
of Whittington and O’Brien usually yielded below PI in the first two years
of the experiment, there may have been extra reserves of nutrients in the
soil to provide the high yields of the third year. It must also be realized
that the apparent degrees of overyielding of several mixtures in a diallel
may be simultaneously affected if, owing to a chance effect, the monocul-
ture yield of one high-yielding genotype is considerably lower (or higher)
than its potential as expressed in other plots. Since the measures of over-
yielding of mixtures in this diallel were not independent at any one harvest
(or between harvests), the multiple instances of overyielding reported by
Whittington and O’Brien do not provide any especially strong grounds for
believing that the overyielding was a real effect.
The second case in which overyielding achieved significance ( P << 0.01
in two independent comparisons) was reported by Rhodes ( 1968b). In
a diallel of ryegrass varieties, under two regimes of cutting (high and inter-
mediate frequency), a particular mixture overyielded by 12% and 15%.
In the infrequent-cutting treatment, this mixture’s yield was below P. The
plots with the different cutting regimes were independent, and so there
seems to be reason to accept this result as due to a real effect. The third
case of significant overyielding was also reported by Rhodes (1970a) in
two mixtures of ryegrass genotypes grown under near-optimal soil condi-
tions. These mixtures overyielded by 11% ( P < 0.01) and 9%
(P < 0.05) under a regime of infrequent cutting; with frequent cutting,
the same mixtures did not yield transgressively.
182 B. R. TRENBATH
A fourth instance of “significant” overyielding has been reported which,

although lacking clear statistical treatment, seems also to be due to a real
effect. Data of Khan (in Harper, 1965) and data of an independent experi-
ment by Harper (1965) both show that mixtures of various flax and/or
linseed varieties overyielded. Khan’s data show two flax-linseed mixtures
as overyielding (significantly,” Harper, 1965, p. 471) by 13% and 14%.
Harper’s own experiment included the same two mixtures. At low density,
one overyielded by 38% while the yield of the other was nontransgressive;
at high density, neither mixture yielded transgressively. Since, at low den-
sity, only two plants were present in each pot, the experimental errors must
have been very large. Furthermore, Harper mentioned neither replication
nor significance levels, and so the value of this experiment as an indepen-
dent confirmation of Khan’s results is uncertain. If Harper’s experiment
was in fact adequately replicated, the high proportion of overyielding mix-
tures (40% ) would suggest that his mixtures were worth further investiga-
tion. While not based on the same varieties, data of Obeid and Harper
(in Harper, 1968) again show a flax-linseed mixture (planted this time
with a range of proportions) which overyielded by a maximum of l o % ,
29%, and 31% at the three densities studied. The flax-linseed
(WEIRA-VALUTA) mixture of Khan remains, however, the only mixture
which has, to this author’s knowledge, ever been reported to overyield in
two separate experimenkl
Little consideration has been given by authors to the possibility that
mixtures may underyield. No cases of significant underyielding appear to
have been reported, although Ahlgren and Aamodt (1939) found that all
three mixtures in a incomplete diallel of grass species underyielded (non-
significantly, by 11 % , 23 %, and 26% ) . Donald ( 1946) repeated this ex-
periment using the same species, but none of the mixtures underyielded.
As mentioned previously, Donald (1963) and van den Bergh (1968)
have each concluded that there is no firm evidence that a mixture can have
an advantage over the higher-yielding component monoculture (mixtures
of legume and nonlegume are excepted), Also, Woodford (1 966) has ex-
pressed the same opinion. The additional evidence presented in this review
is consistent with these conclusions. However, it seems that “firm evidence”
has rarely been sought. Experiments showing transgressive yielding by mix-
tures have either seldom been repeated, or if they have, the results have
not been published. Until the transgressive yielding of more mixtures can
be shown to be repeatable, the addition of further data concerning once-
‘There are only two analogous cases in the production of grain. One is the
mixture of rice varieties (BK and 2A) of Roy (1960), and the other is a mixture
of wheat varieties (RAMONA and BAART) of Chapman ef al. (1969) and Allard and
Adams (1969).
performed experiments can add little to what is already known about mix-
ture performance.
111. Theoretical Considerations
Let us assume that a 1 : l mixture of the seeds of two genotypes has

been sown along with monocultures of the components. If plants of each
component yield the same as they do in their respective monoculture, the
yield of the mixture will be the mid-monoculture yield ( P ) . The plants
of the components of the mixture will be said to have given their “expected
yields” (Alcock and Morgan, 1966) since they have yielded in accordance
with their genotypic potential, as expressed in monoculture. The mid-
monoculture yield will be the corresponding “expected yield” of the
mixture.
Writing the per-plant yield of genotype i in mixture with genotype j as
Y i j ,the average per-plant yield, M i j , of the i, j mixture will be:
Mij =
1
-(Yij
a
+ Yji)
If per-plant yields of i and j in the mixture are the same as in their

monocultures, i.e., if Y i j = Yii and Y j i = Y j j , the mixture yield is the
“expected yield,” i.e., M =: P. In such a mixture, the average per-plant
yield would be given by
Mij =
1
-(Yii
a
+ Y,jj)
If the density of plants is n plants per plot of unit area, then M ,P,,and
P, (the yields per unit area of the mixture and of the higher and lower
yielding monocultures, respectively) are given by
M = nM;,
PI = nYii (for Yii > Yjj)
Pz = nYjj (for Y ; ; > Yjj)
However, as many early experiments showed (e.g., Montgomery, 1912;

Tansley, 191 7; Clements and Weaver, 1924; Sukatschew, 1928), per-plant
yields of genotypes in mixture and monoculture are seldom the same. From
this springs the tendency for mixture yields to deviate from the “expected
yield.”
The biological processes responsible for the deviation of component per-
formance from that expected are complex and varied. A general term which
has been applied to them is “interference” (Crombie, 1947; Harper,
184 B. R. TRENBATH
1961). They result in what may be called either “interference effects” or

“neighbor effects” (“Nachbarwirkungen,” Lampeter, 1960). The latter is
a more objective and neutral term and, hence, perhaps preferable (Tren-
bath and Harper, 1973).
The best understood, and perhaps the most important, mechanism that
can cause the biomasses of plants of a genotype to differ between mixture
and monoculture is the process of “competitionyy(Clements et al., 1929;
de Wit, 1960; Donald, 1963). Plants are conceived as “competing” for
the limited supplies of environmental resources necessary for growth. Clem-
ents (1904) showed how the anthropomorphic overtones of this concept
could be avoided when he asserted that the reaction of a plant to neighbor-
ing individuals is not a direct response to the neighbors themselves, but
to the plant’s own microenvironment insofar as it has been altered by the
presence of the neighbors. In a mixture, differences of morphology and
physiology between the mixture components cause their individuals to
experience different microenvironments and hence different resource
availabilities from those experienced by plants of the same genotype in
monoculture. As a consequence, resources are unevenly shared between the
components and per-plant mixture yields deviate from “expected yields.”
Such effects are said to be due to competition.
De Wit ( 1960) has presented a model of intergenotypic competition
based on the simple assumption that the biomass yield of each component
is strictly proportional to the share of environmental resources it can ac-
quire, According to this model, if the sharing is uneven, plants of one geno-
type, say i, will be larger in mixture than in monoculture while plants of
the other component, genotype 1, will be correspondingly smaller. In such
a case, genotype i is termed the “aggressor” (Donald, 1946) and genotype
j may be termed the “subordinate.” Hence, according to this simple model
of unequal sharing of environmental resources, Yij > Yii and
Yji < Y j j .This appears to be the commonest situation in mixtures, for
of 70 results reviewed by Donald* (1963), 51 (74%) were of this type.
Among the 344 results reviewed in Section 11, there are 326 cases where
data are available of the performance of the components within the mix-
tures; of these 326 mixtures, 255 (78%) showed the same pattern. The
more specific predictions of the de Wit (1960) model have also been
shown to be fulfilled in many field and pot experiments using gramineous
species (de Wit, 1960; van den Bergh, 1968).
When the per-plant yield of one genotype is higher in mixture than in
monoculture and the per-plant yield of the other genotype is correspond-
ingly lower, then the behavior of the mixture components is said to be
Of the 70 mixtures, 34 were of a grass and a legume; in 10 mixtures the density
was not closely controlled.
of a “compensating” type (Aberg et al., 1943; Donald, 1963). If the plant

relative yield (PRY) (based on the per area relative yield of de Wit and
van den Bergh, 1965) of a component is defined as the ratio of the per-
plant yield in mixture to that in monoculture, then in such a mixture, the
PRY of the aggressor will be greater than unity; that of the subordinate
will be less than unity. Thus, if genotype i is the aggressor, Yii/Yi,i> 1
and Y i i / Y i j < 1.
The terminology of this relationship has unfortunately been confused
by Schutz and Brim (1967), who applied the term “complementary” to
mixtures in which, as above, deviations of PRY from unity are of the type
(+,-) . “Complementary” as applied to mixture components had already
been used in the botanical literature to refer to something rather different
(Salisbury, 1929; and see Section V ) . Among other terms which they intro-
duced, Schutz and Brim used “neutral” for cases where both components
give their “expected yields,” i.e., cases of the type (0,O). Mixtures of the
type (+,O) and (-70) were described by Schutz and Brim as showing
“over-compensation” and “under-compensation,” respectively. While “neu-
tral” and “over-compensation” and “under-compensation” seem to be use-
ful terms, the original term “compensating” (noun “compensation”) will
here be retained for cases of the type (+,-) .
With respect to mixture productivity, the common occurrence of com-
pensation tends to keep mixture yields between the monoculture values.
According to the de Wit (1960) model, when components in a 1 : 1 mixture
are competing for the same supplies of environmental resources, the pro-
portional increase of per-plant yield of one component will tend to equal
the proportional decrease of per-plant yield of the other (see Fig. 1) . This
implies that the mean of the PRY’S will have a value close to unity. In
terms of the per-area relative yields of de Wit and van den Bergh ( 19651,
this value is a total which they have called the relative yield total (RYT).
The RYT is given in the present notation by
-+-
R Y T = -1 (Yii G;)
2 Yii
Van den Bergh ( 1 968) showed that if RYT = 1, the mixture yield must
lie between the yields of the pure cultures (strictly P , 2 M 2 P 2 ) . The
general scatter of observed RYT’s around the value of unity is shown for
572 mixtures’in Table 11. These data seem to provide ample basis for ex-
pecting RYT’s to lie close to unity, and incidentally provide support for
the wide applicability of de Wit’s (1960) model. The asymmetry of the
low and the high deviations from unity (13.6% compared with 20.3%)
is significant at the 1% probability level.
186 B. R. TRENBATH
Quontity of resource acquired / plant
FIG. 1. Graphical interpretation of de Wit's (1960) model when components of

a 1:l mixture are competing for the same environmental resource. Compared with
the unit quantity of resource acquired in monoculture, each plant of the aggressor
gains an extra amount, F, in mixture. The quantity of resource acquired by each
plant of the subordinate is correspondingly reduced by F in mixture. The per-plant
acquisition of resource is assumed to be the s_ame in the two monocultures. M,,
is the average per-plant yield of the mixture; P/rt is the mid-monoculture yield in
per-plant terms; Y,,(open circle) and Y i , (filled circle) are per-plant yields of
genotype i in monoculture and mixture respectively; Y,,(open square) and Yji (filled
square) are corresponding per-plant yields of genotype j . Arrows by the monoculture
points lie in the direction of the points of the same genotype grown in the mixture.
The numerical values shown for biomasses indicate that in such a system, the
proportional increase of Yl, over Yii equals the proportional decrease of Y , c
below Y j , .
IV. Types of Interaction Causing Nontransgressive Deviations

of Mixture Yields from Mid-Monoculture Values
Since the concept of relative yield total (RYT) has been introduced
in the preceding section, it can be immediately explained that the present
section treats only those mechanisms (chiefly competition) which seem
able to cause deviations of M from I' without any deviation of RYT from
unity. Such deviations of mixture yield are necessarily nontransgressive.
(Other mechanisms of interaction exist which can cause nontransgressive
deviations from I' but with RYT not equal to unity. Since these mecha-
nisms are also potentially capable of giving rise to more extreme deviations,
they are treated in Section V.) We examine first the nature of each mecha-
TABLE I1
Distribution of the Relative Yield Totals of Mixtures Based on
Published Data of Biomass of Components in 572 Mixtures
~ ~~~
RYT value
0.5 to 0.7 to 0.9 to 1.1 to 1.3 to 1.5 to

Crop 0.7 0.9 1.1 1.3 1.5 1.7 21.7 Author
Grasses 1 2 Ahlgen and

Aamodt (1939)
Either grasses 1 4 1 Aberg et al. (1943)
or legumesb 3 9 2 Donald (1946)
Wheat 2 1 Sakai (1953)
Rice 6 26 4 Sakai (1955)
Grasses" 5 41 11 1 Lampeter (1960)
A series of
nonlegumesb 4 8 3 Williams (1962)
Subterranean
clover 3 Williams (1963)
Flax and
linseed 2 5 5 9 3 3 3 Harper (1965)
Grassesa 3 95 22 England (1965)
Barley 10 Norrington-
Davies (1967)
G rassesa 7 20 9 ' L - 2 Norrington-
Davies (1968)
Grasses. 3 106 4 van den Bergh
(1968)
Grassesa 3 21 26 21 4 Whittington and
O'Brien (1968)
Grasses. 6 5 2 3 Tbomson (1969)
Rye 6 18 6 Norrington-
Dnvies and
- - - - - Hutto (1972)
6 72 378 95 13 5 3 = 572
1.0% 12.6% 66.1% 16.6% 2 . 3 % 0.9% 0.5%
13.6% 20.3%
~~ . . . . . . . . . . . . . . . . . . . . . . . . .
Data derived from a series of cuts.

Mixtures of leguminous and nonleguminous species have been omitted.
nism and then consider how it may influence the relationship between the
yield of a mixture and the yields of its component monocultures.
The environmental resources for which plants compete are principally
the light, water, and soil-nutrient supplies necessary for growth (Clements
et al., 1929; Harper, 1961; Donald, 1963; Risser, 1969; Rhodes, 1970b).
Although carbon dioxide is required for shoot photosynthesis and oxygen
188 B. R. TRENBATH
is needed for the respiration of the roots, competition for these seems nor-
mally unlikely to occur. Competition for C02 is probably absent because
the air within the canopy is generally well mixed (Monteith, 1963; Impens
et al., 1967) ; all leaves are surrounded by air of approximately the same
CO, concentration. Similarly, except in almost waterlogged soils, the diffu-
sion of oxygen in soil is usually fast enough to maintain adequate supplies
to all roots (Greenwood, 1969).
Apart from the relatively small number of instances where chemical se-
cretions by one species are thought to have influenced the growth of
another, it seems that most investigators who have reported results of mix-
ture experiments in English-language journals appear satisfied that the
neighbor effects which they observed were due to competition for light,
water, or one of the major nutrients (nitrogen, phosphate, and potassium),
or a combination of these factors. Since it is relatively easy to measure
and explain the unevenness of the sharing of light between the foliage of
two components in a mixture, the studies involving competition for light
have generally been the most conclusive.
Clements and many others have related differences in success in mixture
to differences in height between the leaves of the components of mixtures,
simply inferring that leaves of the shorter component must be experiencing
some degree of harmful shading. More recent analytical approaches have
been based on methods suggested by Monsi and Saeki (1953). Working
with natural communities, Monsi and Saeki introduced techniques for the
measurement of profiles of light intensity and leaf area index (LAI) . Since
then, these techniques have been used very successfully in experimental
mixtures to relate the differences in growth rates of the two components
to differences in the proportions of the total incident light intercepted by
leaves of the two components (Black, 1958; Iwaki, 1959; Stern and
Donald, 1962a,b; Williams, 1963). The general conclusion from all experi-
ments involving competition for light is that the component with its leaf
area higher in the canopy is at an advantage. It is also likely (Stern and
Donald, 1962a) that, if the leaves are horizontal, the advantage is greater
than if they are erect (this is because horizontal leaves intercept more of
the total downward light flux per unit area of leaf than do erect leaves).
If the taller component has a greater leaf area, its advantage is again corre-
spondingly greater (Iwaki, 1959).
In most of the experiments cited above, the investigators have attempted
to exclude the possibility of competition for soil factors. By providing opti-
mal soil conditions or by separating the roots of the mixture components,
it was intended that neighbor effects would be interpretable in terms of
competition only for light. In most agricultural environments, however,
soil conditions are suboptimal and since root systems usually interpenetrate
each other freely, there is a possibility of competition for water and/or

nutrients. Where the soil is very infertile and the density of planting low,
competition between root systems for soil factors is likely to decide which
component becomes the aggressor because the leaf area produced may
never become great enough to lead to significant competition for light. In-
deed, large neighbor effects have been reported in cases where shading
was claimed to have been absent (Pavlychenko and Harrington, 1935;
Myers and Lipsett, 1958). In experiments in containers where competition
for light has been prevented by partitions, competition for soil factors has
similarly been shown to produce large effects (Donald, 1958; Aspinall,
1960; Rhodes, 1 9 6 8 ~Snaydon,
; 1971; Eagles, 1972).
The principles involved in competition between root systems have not
been as well worked out as in the case of competition between shoots,
but Bray (1954) has outlined a general theoretical approach. In contrast
with supplies of light, nutrient supplies are usually not greatly added to
after the beginning of growth of an annual crop although subsequent rain-
fali or irrigation normally supplements the initial store of soil water.
Whereas success of a species in competition for light depends on it having
a large absorptive area closer to the light source than the leaves of the
other component, the variable and complex geometry of root systems and
the sources of water and nutrients makes it less easy to define the plant
characteristics likely to confer competitive success.
Bray (1954) noted that competition between root systems for nitrogen
is likely to start at lower root densities, i.e. earlier in growth, than competi-
tion for phosphorus or potassium. This is because nitrogen is much more
mobile in soil than phosphorus and potassium. Zones of nitrogen depletion
round individual roots will therefore begin to overlap relatively early. To
test this proposition, Andrews and Newman (1970) grew pure and mixed
communities of young wheat plants in soil deficient in nitrogen and phos-
phorus with root densities between 1.2 and 8 cm/cm3; their resuIts showed
that whereas competition for nitrogen was intense, competition for phos-
phorus was indeed slight. Since the mobilities of nitrogen and water in
soil are similar (Fried and Broeshart, 1967), the soil resources most likely
to be subject to competition are nitrogen and/or water.
If Bray’s propositions are correct, they allow the identification of the
characteristics that will determine the relative aggressiveness of the compo-
nents of a crop mixture growing on infertile soil. For one component to
gain an advantage over the other in the early competition for nitrogen and
water, a faster growing root system (i.e., generally a greater length of root)
is required. The limited evidence so far available does indeed suggest a
possible correlation between seedling competitive ability and early root pro-
duction (see review in Rhodes, 1970b). However, it also seems clear that
190 B. R. TRENBATH
if root lengths are similar, the genotype having the more widely spreading,
less-branched root system will be at an advantage. Theoretical consider-
ations suggest that there will probably also be an advantage in having roots
as thin as possible, thus allowing the available root material to be present
as the maximum length of root (Olsen et al., 1962). Abundant, long root
hairs (see Olsen and Kemper, 1968) and a high root “demand” factor
(Drew et al., 1969) are also likely to contribute to competitive success.
This theoretical consideration of competition for soil factors has treated
competition between root systems (root competition) as independent of
competition for light between shoots (shoot competition). However,
Donald (1958) has suggested that in well developed agricultural crops of
uniform genotype, both shoot and root competition are usually occurring.
The relative importance and time of onset of root and shoot competition
will depend on environmental conditions (Aspinall, 1960) ; the nature of
the genotypes involved will also have an effect. To compare, for a particu-
lar set of conditions and genotypes, the contributions of the effects of root
and shoot competition with the overall effect of being grown in mixture,
pot experiments have been designed in which partitions separated either
roots or shoots, or both, or none (Donald, 1958; Griimmer, 1958; Aspinall,
1960; Rhodes, 1968c; Snaydon, 1971; Eagles, 1972).
Of the results of these experiments, those of Snaydon (1971 ) are the
easiest to interpret, for in his experiment alone were there constant
numbers of partitions per treatment and plants per compartment. Using
ecotypes of white clover, Snaydon confirmed the results of the other inves-
tigators who had shown that root competition had a greater effect (and prob-
ably began earlier) than shoot competition. Although this finding suggests
that dominance-suppression relationships in crop mixtures may depend
more on root characters than on shoot characters, the concentration of
roots at the surfaces of the containers in such experiments must have
hastened the overlapping of zones of depletion around individual roots,
and hence led to an overestimation of the agronomic effect of competition
for soil factors.
Considering a possible interaction between the two types of Competition,
Donald (1958) showed that the proportional reduction in the subordinate’s
yield due to shoot competition was greater when root competition was oc-
curring than when it was absent. Similarly, the proportional reduction due
to root competition was greater when shoots were competing for light than
when they were separated. Donald suggested that the failure of, say, the
root to acquire sufficient nitrogen caused leaf development to suffer. This
in turn reduced the supplies of assimilate to the roots which grew less and
so were less able to compete for nitrogen. Donald proposed a similar set
of causes and effects for a component of a mixture which was unsuccessful
in competition for light. The important conclusion was that the effects of
Competition, once initiated, tend to be magnified by a system of positive
feedbacks if simultaneous shoot and root competition are occurring.
Milthorpe (1961) has discussed the effects of the onset of competition
for soil factors in a mixture where competition for light has already caused
some depression of the growth of one component. Arguing from the depres-
sive effect of shading on root:shoot ratio (e.g., Brouwer, 1966), Milthorpe
considered that any deficiency of soil water or nutrients would cause the
accelerated suppression of the subordinate if any previous shading had re-
duced its root’s uptake capacity relative to the size of its shoot. Similarly,
the writer has presented results (Trenbath and Harper, 1973) which sug-
gest that the development of a slight deficiency of soil factors where com-
petition for light was already occurring caused an approximately 4-fold in-
crease in the depression shown by a series of subordinates.
While competition for resources has usually seemed able to explain ob-
served differences between per-plant yields in mixture and monoculture,
a number of workers believe that these differences (neighbor effects) are
sometimes caused by other processes. This belief has been expressed by
workers well acquainted with the mechanisms of competition (Warne,
1953), has been implied by other investigators (Sakai, 1955), and has
been used as a convenient explanation of unexpected results by others
(Ahlgren and Aamodt, 1939; Went, 1942).
As an alternative explanation of neighbor effects, allelopathy (Grummer,
1955) has been frequently proposed. The type of allelopathy most likely
to occur in a field crop is that described by Winter ( 1961), where one
or more biologically active substances are liberated from the root or shoot
of a plant, enter a neighboring plant, and there cause a depression of that
plant’s growth. T o allow for cases of stimulation of growth in the affected
plant, the general term “allelochemical” effect (Anonymous, 1971 ) has been
introduced to include both positive and negative effects on growth. Since,
however, the term “allelopathy” is both more convenient and more famil-
iar, in what follows it will be used in a general sense, i.e., allowing the
possibility of either positive or negative effects.
Allelopathy in plant communities has been studied most actively in the
Soviet Union and Germany; the rapidly expanding literature has been re-
viewed by Griimmer (1955), Rademacher (1959), Grodzinsky (1965,
1971), and Borner (1971). On the other hand, in some other European
countries and in the United States, research on allelopathy has had an un-
even history. Several instances of apparent allelopathy have been shown
(sometimes by the original investigators) either to be due to other causes
(Went, 1942; Gray and Bonner, 1948; but see Muller, 1953; Grummer,
1958; but see de Wit, 1960; Sandfaer, 1968; but see Sandfaer, 1970a,b)
192 B. R. TRENBATH
or to be explained by alternative mechanisms (Muller, 1966, but see Bar-

tholomew, 1970). These cases have tended to make investigators cautious
of suggesting allelopathy as the cause of neighbor effects, indeed, they have
inclined the present author to agree with Harper’s (1965) opinion that
research into the role of allelopathic substances in mixtures is beset with ex-
treme technical difficulties. To avoid greatly extending this review no detailed
discussion will be attempted, but three general comments will be made.
First, as Borner (1960) pointed out, although many investigators have
demonstrated allelopathic effects between plants growing in solution culture
or sand culture, instances where the same effects have been conclusively
proved to be significant under field conditions are rather few. Among such
cases reported in English-language and western European literature, the
most convincing seem to be the existence of inhibitory effects on associated
plants due to walnut (Massey, 1925; Bode, 1958) and to Arctostaphylos
species (Hanawalt, 1971) . Also, the demonstration of autotoxicity among
individuals of Grevillea robusta (Webb et al., 1967) grown in both the
field and glasshouse seems conclusive. It should be noted that although
Bonner (1946) demonstrated the autotoxic activity of root exudates of
guayule plants in gravel culture, the active substance was degraded so fast
in field soil that no autotoxic effect was likely under normal agricultural
conditions. A rather different case is that of Hirano and Kira (1965), who
demonstrated apparent autotoxicity among densely planted peach saplings
in an experiment planted in field soil (clay loam) under glass; this result
accorded well with expectations (Hirano and Morioka, 1964; Proebsting
and Gilmore, 1940). However, a similar experiment carried out on a sandy
soil in the open gave no indication of an autotoxicity effect (Hirano and
Kira, 1965).
Second, since crops are normally grown at densities high enough for
competition between neighbors for resources to be intense (Donald, 1963) ,
any allelopathic effect would be either exaggerated (or reversed) through
competition. To emphasize the importance of taking into account both as-
pects of the total interaction, we may consider an hypothetical example
of a crop mixture growing on a soil poor in nitrogen. If the suppression
of one of two otherwise evenly-matched components was initiated through
some slight allelopathic inhibition of its nitrogen uptake, later competition
for nutrients might result in plants of this component being nitrogen defi-
cient and heavily suppressed. The agronomist would “prove” the suppres-
sion to be due to competition for nitrogen since he would find, in an experi-
ment with separated shoot systems, that neighbor effects were strongest
on soils with the lowest nitrogen level; in the meanwhile, the specialized
investigator of allelopathic effects might have discounted some slight inhibi-
tory effect as insignificant in view of the great intensity of the suppression
in the field. Through the lack of an integrated approach, both workers
would have drawn false conclusions. The experimental separation of the

effects of competition and allelopathy is thus an important objective, al-
though it is certain to be difficult to achieve (Welbank, 196l ) .
Third, providing (a) that an allelopathic inhibition takes effect early
enough in the life of the crop for compensatory growth to occur in the
other component (see Section V ) , and (b) that the inhibition reduces the
uptake of growth factors rather than the efficiency with which they are
used, the total quantity of environmental resources intercepted by the mix-
ture may not be much affected. Consequently, the yield may scarcely be
less than that of another mixture with the same values of P, and P , in
which a similar degree of suppression has been brought about by competi-
tion acting alone. Where, however, the allelopathic effect reduces the effi-
ciency with which growth factors are used in dry-matter production, the
RYT will probably be lowered and the mixture could underyield (see Sec-
tion V) .
Whether caused by competition alone or by a combination of competi-
tion and allelopathic effect, the unequal sharing of resources between com-
ponents generally affects the yields of mixtures. If the higher-yielding com-
ponent in monoculture is the aggressor, then with RYT = 1 the mixture
yield will lie between f' and P I . The greater the depression of the lower-
yielding component, the closer the approach of M to P,; with its complete
suppression, M = PI. Similarly, if the lower-yielding component in mono-
culture is the aggressor in the mixture (the so-called Montgomery effect,
Gustaffson, 1951), then M lies between P and P,. With complete suppres-
sion of the subordinate, M = P,.
The tendency for mixture yields to lie above P (Table I) combined with
the closeness of RYT's to unity (Table 11) suggests that there is a positive
correlation between aggressiveness in mixtures and biomass production in
pure stands. The association is not strong, however, for the value of the
correlation coefficient has been estimated from published data to be only
about 0.3 (Trenbath, 1972). The existence of a positive correlation is
nevertheless in agreement with the ideas discussed earlier, namely, that
a large leaf area displayed at a sufficient height gives an advantage in com-
petition for light (e.g., Donald, 1961) ; the close dependence of productiv;
ity in monoculture on LA1 is well established (Watson, 1952).
A possible reason for the looseness of the correlation between aggres-
siveness and monoculture yield is found in a suggestion by Iwaki (1959).
Iwaki postulated that a species would be aggressive in mixture if it diverted
a particularly large share of photosynthetic product into building tall stems.
Associated species would be shaded and suppressed. In monoculture, how-
ever, the tall species would grow relatively slowly because of the low pro-
portion of its dry matter invested in productive leaves. A negative correla-
tion between aggressiveness and monoculture grain yield in rice (Jennings
194 B. R. TRENBATH
and Aquino, 1968) was similarly related to the tallness of the stems of
one type of rice, but in this case the low monoculture yield of the tall
type was mainly the result of its tendency to lodge. The bending over of
the tips of long grass leaves (“flagging”) will also lead to greater aggres-
siveness (see above) and lower growth rates in monoculture (Alberda,
1966).
The aggressiveness of a species is well known to depend on environ-
mental conditions and such a dependence will affect the relative values
of M ,P I , and P,. The dominance-suppression relationship between geno-
types has been reversed by changing the temperature regime (Eagles,
1972) and by aItering the soil conditions (van Dobben, 1955; Sakai and
Iyama, 1959; Stern and Donald, 1962a; van den Bergh and Elberse, 1962;
Snaydon, 1971). Aggressiveness often depends on the stage of growth of
the plants (e.g., de Wit and van den Bergh, 1965; van deli Bergh, 1968;
Rhodes, 1968a; Nguyen Van, 1968), but in field experiments, the concur-
rent changes in developmental stage and in meteorological factors usually
make it difficult to identify the factor responsible for any change of aggres-
siveness with time. The hard-to-define differences between growing seasons
have marked effects on aggressiveness at least where the index used to
measure it is based on seed yields (Laude and Swanson, 1942; Sakai and
Oka, 1955; Allard and Workman, 1963; Workman and Allard, 1964; Lin
and Torrie, 1968). Allard et al. (in Edwards and Allard, 1963) showed
from the data of Suneson (1949) that taking the value of ATLAS as 100,
the selective advantage of VAUGHN ranged from 40 to 173 in 13 seasons.
Transfer from the field to the greenhouse may also cause reversals of domi-
nance (Aberg e f al., 1943; Syme and Bremner, 1968).
If aggressiveness is so sensitive to environmental conditions, it might
be asked how this sensitivity affects mixture yields over a series of sites
or seasons. Unfortunately, although diallel experiments have been carried
out using a range of environments or treatments (Harper, 1965; England;
1965; Whitehouse et al., 1967; Bell et al., 1968; Norrington-Davies, 1968;
Sandfaer, 1970b; Schutz and Brim, 1971), no systematic attempt has yet
been made to study the effect of associated changes of aggressiveness on
mixture yields. Presumably connected in some way with such changes, a
greater stability of seed yields has often been found in varietal mixtures
of cereals and soybeans grown over a range of environments [Allard, 1961;
Simmonds, 1962; Pfahler, 1965; Frey and Maldonado, 1967; Byth and
Weber, 1968; Qualset and Granger, 1970; Schutz and Brim, 1971; but
see Rasmusson (1968) and Clay and AIIard (1969)l. Taking together
the results of three reports (Allard, 1961; Pfahler, 1965; Qualset and
Granger, 1970) in which an index of yield stability was given for each
mixture and monoculture, 5 out of a total of 12 binary mixtures were more
stable than their more stable component monoculture; the remaining mix-
tures showed stabilities between those of their component monocultures.
In a comprehensive study involving 16 environments, Schutz and Brim
(1971) calculated, for each type of stand, indices of stability with respect
to several types of environmental variation (seasons, locations, replicates,
etc.) . The lack of agreement between the several stability indices of individ-
ual types of stand suggests a complex situation with no clear-cut stability
advantage for mixtures. To add to difficulties of interpretation, stability
parameters are much less accurately estimated than parameters related di-
rectly to yield (Allard, 1961;Marshall and Brown, 1973) .
With reference to biomass rather than to seed yield, very few data are
available that allow the comparison of any index of yield stability of indi-
vidual mixtures with those of their component monocultures. While two
studies (England, 1968; Sechler and Chapman, 1967) suggested that mix-
ture yields were collectively more stable than monoculture yields, data of
individual mixtures from two other studies (Pfahler, 1965; Thomson,
1969) showed stabilities to be intermediate between those of the monocul-
tures in 9 out of a total of 10 cases. These limited data seem to indicate
that the stability of mixture yields is, like yield itself, usually nontransgres-
sive, and that in seed yield, but not in biomass yield, there is a tendency
for mixture stability to be close to that of the more stable component.
In a theoretical study of mixture stability, Marshall and Brown (1973)
suggested that varietal mixtures were most likely to show agronomically
useful stability in highly variable environments to which the available geno-
types were not individually well adapted. Perhaps it is partly the lack of
crop species with sufficiently wide adaptation that has led to the extensive
culture of multispecific mixtures in India (Aiyer, 1949). The markedly
differing adaptation characteristics of the components of such mixtures en-
sure that even in the worst season there will be something to harvest.
The principal conclusions from this and the previous sections may now
be summarized. In field-crop mixtures, competition for both light and soil
resources will usually be occurring; allelopathic effects (if present) will
operate in conjunction with this competition; competition for resources
and/or allelopathy will usually cause per-plant yields of each genotype in
mixture to differ from that in monoculture; the differences of per-plant
yields are usually of a compensating type; if RYT = 1, M will lie between
PIand P,;any positive correlation between aggressiveness and monoculture
yield will produce a tendency for M > P; there are some indications that
the biomass yield of a mixture having sufficiently different components is
likely to be more stable than that of the more stable of its components
in monoculture. We continue now by turning to the more varied and subtle
forms of interaction that may lead to transgressive yielding.
196 B. R. TRENBATH
V. Mechanisms Capable of Causing Transgressive Yielding by Mixtures
The results reviewed in Section I1 show that mixtures have often been
recorded as apparently yielding transgressively. Furthermore, the data indi-
cate that records of mixtures overyielding are significantly more frequent
than records of underyielding. Two contrasting interpretations of this situ-
ation could be proposed.
1 . It might be suggested that if RYT could be measured without experi-
mental error, its value might always be close to unity; the observed skew-
ness of its distribution (Table 11) could be a consequence of basing RYT
on two ratios of random normal variables (Fieller, 1932). Since there is
a weak, positive correlation between aggressiveness and monoculture yield
(Section IV) , the preponderance of mixture yields greater than P could
be the result of this correlation. If this were the case, theoretical error-free
experiments would show mixture yields lying usually between P, and p,
less frequently between P and P,, and never outside the range P, to P2.
The experimental error found in actual experiments would be expected
to disperse the observed results about their “true” values, so that overyield-
ing mixtures would be recorded more frequently than underyielding ones.
This view implies that observed cases of transgressive yielding are due only
to experimental error.
2. An alternative interpretation might be that at least some of the ob-
served cases of transgressive yielding do represent real effects. If the corre-
lation between aggressiveness and monoculture yield were discounted as
being too weak to have appreciable effect, the tendency for mixture yields
to exceed P could be attributed to apparent mutual stimulation of the mix-
ture components, as indicated by the preponderance of RYT’s greater than
unity (Table 11).
These two interpretations are only preliminary attempts to explain the
observations; they represent extreme, but not necessarily incompatible, ap-
proaches. Having discussed in Section I1 the evidence in favor of transgres-
sive yielding being a real phenomenon, we consider here the mechanisms
that could lead to transgressive yielding. If they appear likely to operate
under the conditions normally used for mixture experiments, this would add
credibility to the second interpretation given above.
In connection with overyielding, the findings and theory of animal
ecology may be relevant since they help to define conditions that might
lead to this kind of transgressive yielding. Gause (1934) quoted Formo-
zov’s observations of natural mixed populations of tern in which four spe-
cies coexisted, apparently because they exploited the environment in differ-
ent ways. In such a case, it seems likely that a mixed population would
be more productive than any monospecific population. Gause (1934)

summed up his own experimental results in the principle that species of
very similar requirements tend not to coexist. Using this principle, Harper
(1967) and others argued that the species diversity found in natural com-
munities implies that the component species occupy differing niches. To-
gether, therefore, they must exploit the environment more completely than
a community of few species. Providing support for this view, Beardmore
et ul. (1960) and Seaton and Antonovics (1967) reported overyielding
of mixtures of Drosophilu strains in controlled environments. Seaton and
Antonovics suggested that in their mixtures the two genotypes were avoid-
ing competition by occupying different niches.
In the botanical literature, genotypes believed to be avoiding competition
while they share a habitat are said to be “complementary” (Woodhead,
in Salisbury, 1929). Woodhead first used the term in connection with natu-
ral woodlands where, by the sharing of environmental resources in either
time or space, the various types of species seem to escape, at least partially,
the effects of competition. His concept of the complementary use of re-
sources would seem to be applicable to crop mixtures; the possible ways
in which resources may be used in such a manner will now be considered.
Van den Bergh and de Wit (1960) reported an example where temporal
sharing of the environment may have been responsible for a case of appar-
ent mutual stimulation in mixture. In a mixture of two grass species which
differed markedly in time of development, plants of both components had
more tillers (53% and 3 6 % ) than did plants in the corresponding mono-
cultures; although biomasses were not reported, it may be noted that the
RYT based on tiller number was 1.49. Syme and Bremner ( 1968) reported
a series of experiments involving oats and barley varieties chosen to differ
in flowering time; the results of an experiment performed under glass (data
given in Syme, 1963) showed that in all four oats-barley mixtures, both
components showed higher per-plant shoot weights than in monoculture.
While overyielding of dry matter did not occur, the RYT based on shoot
weights averaged over the four (replicated) mixtures and two densities
was 1.12. Such an RYT value in a mixture of which the two monoculture
yields were very similar would be associated with the mixture outyielding
the monocultures by 12%. Sechler and Chapman (1967) also reported
an experiment with cereal genotypes (oats) which differed greatly in
flowering date. Again no mixture overyielded. Unfortunately, the data
given were insufficient to allow the calculation of RYT’s and so no detailed
comparison is possible with the experiment of Syme and Bremner ( 1968).
In corn-rice mixtures in the Philippines, the corn flowers and matures
before the rice begins to flower; this phenological difference probably ex-
plains the reported overyielding of grain by such a mixture (International
198 B. R. TRENBATH
Rice Research Institute, 1974). Flax and linseed differ in maturity date
and their overyielding of biomass in several mixtures was attributed to this
by Obeid ( 1965) (see Section 11).
Differing temporal patterns of growth or development sometimes result
in a reversal of dominance during the growth of a mixture (Harper and
Clatworthy, 1963; England, 1965). Even if the biomass increments of the
mixture over the periods before and after the reversal are not transgressive
compared with the increments in the monocultures over the same intervals,
the biomass accumulated by the mixture over the whole growing season
may yet be transgressive. Van den Bergh (1968) gave a hypothetical exam-
ple of this in which the more aggressive component in each phase was
the component with the greater biomass-increment in its monoculture dur-
ing that phase. In this example, the total accumulation of biomass over
the two phases was the same in each component monoculture; the associ-
ation of greater aggressiveness and faster biomass accumulation within each
phase resulted in overyielding by the mixture at the final harvest. If the
direction of this association had been reversed, the result would have been
an underyielding mixture. Examples of such effects have not yet been
found, and too few suitable sets of data exist to judge their plausibility.
Van den Bergh (1968), failing to find this kind of effect in the data of
England ( 1965), suggested that the originally suppressed species had been
unable to take advantage of the sharp reduction in the vigor of the other
component of the mixture.
The components of a mixture may be complementary in a spatial sense
by exploiting different layers of the soil with their root systems. Although
Gustaffson’s ( 1954) discussion and vague reference to such a case cannot
be taken as more than an indication, he claims the possibility of increased
grain yields in cereal mixtures through the stratification of root systems.
Cable (1969) found that desert plants were affected least by the proximity
of plants of other species when the root systems of these neighbors did
not overlap the depth of their own. Mutual avoidance^' by adjacent root
systems (Raper and Barber, 1970; Baldwin and Tinker, 1972) could lead
to a late-developing root system occupying deeper soil horizons in mixture
than in monoculture. The significant overyielding of grass mixtures re-
ported by Whittington and O’Brien (1968) was accompanied by phos-
phorus uptake from greater depths in the mixtures than in the monoculture
(O’BrienBet al., 1967) and so might be due to such an effect. However,
the different pattern of uptake in mixture could also have been the result
rather than the cause of enhanced growth. The writer (Trenbath, 1970),
measuring panicle weights in a field experiment involving 5 mixtures of oat
species, found in one replicate that 5 out of 5 mixtures overyielded; in
the succeeding two replicates in a linear sequence of three, the numbers
of overyielding mixtures were, respectively, 2 and 1 out of 5. The reality

of this trend was supported by the significance (P < 0.01) of the regres-
sion of panicle-weight RYT’s of the 5 mixtures on the position of the repli-
cate in the linear sequence across the field. The trend in mixture yields
was tentatively related to an observed soil-depth gradient. It was suggested
that stratification of root systems had occurred on the deep soil, leading
to high RYT’s and overyielding, but that this stratification had been pre-
vented on the shallow soil. The two species making up the mixture with
the highest RYT were suspected to differ the most in depth of rooting.
Ellern et al. (1970) showed that indeed these two species had rooting
depths significantly different; the other species were unfortunately not
tested.
Components of a mixture may complement each other nutritionally; one
component may require much of an element of which the other component
needs little (see Kolb, 1962; Davies and Snaydon, 1973). Considering a
particular element, one component may be able to utilize a form that is un-
available to the other. Although mixtures of leguminous and nonlegumi-
nous species are not formally treated in this review, it is such mixtures
which provide the most striking and repeatable examples of overyielding
due to nutritional complementation. Since the roots of the components are
drawing on different supplies of nitrogen (soil and atmospheric sources),
on nitrogen-deficient soils, RYT values regularly exceed unity and over-
yielding is often recorded (e.g., de Wit et af.,1966; Ennik, 1969). When
nodulation is prevented, the species compete for the same supply of nitro-
gen, the RYT falls to unity, and overyielding does not occur (de Wit et
a!., 1966). Since phosphorus is present in the soil in several forms of differ-
ing availability to different species (Richardson et al., 1931; Schander,
1941 ) , nutritional complementation with respect to phosphorus could also
occur but has not yet been reported.
Overyielding by mixtures has in some instances been attributed to a more
efficient utilization of light by their canopies. The use of mathematical
models has suggested that the highest photosynthetic rate might be obtained
from a canopy in which the steepness of the inclination of the leaves de-
creases with depth (Warren Wilson, 1960; Verhagen et al., 1963; Duncan
et al., 1967; Nilson, 1968). This “ideal” leaf arrangement could be ap-
proached by a mixture of a tall erect-leaved genotype and a short, prostrate-
leaved one. A mixture of such components might possibly overyield. In
addition, Verhagen et al. (1963) have used a simple model to show theo-
retically that to maximize photosynthetic production, as the leaf area of
a crop increases, the average inclination of the leaves should also increase.
I n the “leaf-inclination” mixture just described, the shorter, prostrate-
leaved form would tend to be progressively suppressed through shading by
200 B. R. TRENBATH
the taller form, and hence the average inclination of leaves in the canopy
as a whole would indeed increase.
To examine further the possibility of mixtures overyielding due to favor-
able canopy configurations, the author (Trenbath, 1972) used a computer
model based on that of Duncan et al. (1967) to simulate daily gross photo-
synthesis of mixtures and monocultures of wheat varieties with contrasting
leaf inclinations. In 3 of the 4 sets of conditions used, mixture advantages
in photosynthetic rate were predicted to appear between LA1 2 and 7
(approximately) ; such advantages depended on the latitude, cloud cover,
and planting time being such that most of the incident light arrived from
relatively high elevations. In a mixture with LA1 4, given the radiation
conditions of latitude Oo at the equinox (sunny, with a solar elevation at
noon of 9 0 ” ) , the RYT based on simulated daily photosynthesis was 1.087.
This RYT corresponded to the unreal, but extremely favorable, situation
of a mixture of nonoverlapping, stemless canopies, the upper canopy with
a leaf inclination of 75O and the lower with an inclination of 1 5 O . The
introduction of stems and the inclusion of a moderate degree of overlapping
of the canopies reduced the RYT considerably. If the components were
to have less extremely erectophile and planophile leaf canopies, the RYT
would be reduced still further and any predicted mixture advantage in
photosynthesis would be very small.
Rhodes (1968b) reported the yield of grass in a mixture whose canopy
structure approached Warren Wilson’s ( 1960) “ideal” configuration men-
tioned above. The mixture overyielded significantly in two treatments with
frequent cutting, but not in the treatment where plots were left to grow
until their canopies intercepted 95% of the incident light. It is unlikely
that this overyielding was due to any similarity between the mixture canopy
and Warren Wilson’s (1960) ideal type since, as already mentioned, theo-
retical experiments (Trenbath, 1972) suggest that the advantage of a “leaf
inclination” mixture (erect-leaved canopy above a prostrate-leaved
canopy) will be greatest in stands with LAI’s between 2 and 7. In a similar
way to Rhodes, van den Bergh (1968) and Alcock and Morgan (1966)
found (nonsignificant) overyielding in grass mixtures where repeated cut-
ting probably prevented the LA1 from exceeding 2 for any extended period.
Although van den Bergh appealed to possibilities of better use of light by
the sods of such mixtures, he gave no measurements or theoretical basis
for such an explanation.
Of extreme interest is Rhodes’ (1970a) finding that two mixtures, of
which the components had contrasting leaf inclinations, overyielded signifi-
cantly (see Section 11) under infrequent cutting, whereas under frequent
cutting they yielded between PI and P,. Such observations accord much
better with theory than do Rhodes’ (1968b) results, but it should be noted
BIOMASS PRODUCTIVITY OF MIXTURES 20 1
( a ) that canopy structures were not actually measured in any of these

studies, and ( b ) that other mixtures which contained apparently similar
combinations of tall-erect with short-prostrate forms (two in Rhodes’
1970a experiment and three in his 1968b experiment) did not overyield.
If yields of individual components had been measured in Rhodes’ studies,
it would have been possible to use RYT values to detect and quantify
positive interactions between components in a manner independent of over-
all yields. It is possible that positive interactions occurred in all Rhodes’
“leaf-inclination” mixtures, but that differences of monoculture yields were
sufficient to prevent their expression as overyielding.
Before leaving the topic of light utilization, it is pertinent to point out
that Duncan et al. (1967) simulated the photosynthesis of a canopy of
which the leaves were horizontal above, becoming more inclined with
depth. According to their result, it can be expected that a mixture of a
tall, prostrate-leaved form and a short, erect-leaved form might underyield.
No detailed models of competition for water appear to have been pub-
lished and hence there are no quantitative predictions of the effect of water
shortage on mixture yields. Nevertheless, considering the growth of meso-
phytic crops under conditions of high radiation and water shortage, Aiyer
(1949) has emphasized the beneficial effect of shade trees or other taller-
growing mixture components; also, Baldy (1963) has argued that the
many-layered mixed communities traditionally grown in desert oases (e.g.,
date palm + +
apricot vegetables) may use applied water more efficiently
in biomass production than pure stands. In such oasis communities, the
shading- and windbreak-effect of the upper layer(s) creates a favorable
microclimate for the layer below; the component chosen for each succes-
sively lower layer is more mesophytic and less light-demanding than the
one above. The shortest component (the vegetable crop) escapes suppres-
sion possibly because soil cultivations tend to prevent intermingling of the
root systems. Where the shorter component grows better under shade than
in the open (Aiyer, 1949) and where the taller component benefits from
the presence of the shorter one (Baldy, 1963, p. 339), water-use efficiency
may be superior in the mixture and overyielding of biomass may occur.
To the author’s knowledge, this possibility has not been rigorously tested.
Allelopathic effects can theoretically cause transgressive yielding. If an
allelopathic substance produced by one component affects the growth rate
of the other component by changing only the rate of uptake of some limit-
ing growth factor, the apparent relative competitive abilities of the mixture
components will change but the total quantity of the factor taken up may
not be much different from that in the absence of allelopathy. If this is
so, RYT will be close to unity (Section IV). If, however, the substance
changes the efficiency with which the growth factor is utilized, RYT will
202 B. R . TRENBATH
deviate from unity and transgressive yielding is possible. Roy (1960) dis-
covered a binary mixture of rice varieties which overyielded in grain weight
in several independent trials. By manipulating the cultural conditions, he
found that the growth stimulation appeared to be caused by some agent
carried between components in the irrigation water. Although the impor-
tance of this result ought to have led to an immediate repetition of the
work, no further reports appear to have been published. Allelopathic stim-
ulation has not so far been invoked as an explanation of overyielding in
experiments where the yield was dry matter.
Another way in which allelopathy could lead to overyielding is suggested
by the experience of foresters in New South Wales and Queensland. Webb
et al. (1967) have reported that six rain-forest trees which do not form
natural pure stands show unexpectedly poor growth in commercial mono-
cultures. Detailed experimentation using one of the species (Grevillea ro-
bustu) indicated that the growth of young individuals was inhibited by a
water-soluble substance apparently produced by the roots of adjacent G .
robusta plants. If the same mechanism is causing the autoinhibition of the
other five species, mixtures of them might overyield. It is to be hoped that
experiments will soon be undertaken to confirm and clarify these and other
possible effects of allelopathy on mixture yields.
Mechanical factors could, again theoretically, lead to transgressive yield-
ing by a mixture. For example, let us suppose first that the component
with the potentially higher yield in monoculture is susceptible to lodging,
and second that the other component resists lodging strongly enough to
cause the mixture to stand while the susceptible monoculture lodges. If
the lodged monoculture yields less than the unlodged mixture, the mixture
is expected to overyield. Such a situation is not unlikely since lodging in
mixtures commonly follows the behavior of the more resistant component
(Atkinson, 1900; Tsedik-Tomashevich, 195 1 ; StringEeld, 1959; de Wit,
1960; B. R. Trenbath, unpublished data; but see Probst, 1957; Qualset
and Granger, 1970).
Although very few critical experiments appear to have been performed
to test the point, a consensus of opinion is developing which maintains
that mixtures have a generally greater tolerance of disease and pest attack
(Schwerdtfeger, 1954; Borlaug, 1959; Simmonds, 1962; Browning and
Frey, 1969; Adams et al., 1971; Cherrett et a]., 1971). When a stand
of susceptible plants is “diluted” with resistant plants, the level of infesta-
tion or damage of individual susceptible plants may be reduced. (Suneson,
1960; Browning, 1966). Since the foliage of the resistant plants acts as
a spore trap, the growth rate of rust in an epidemic is reduced (Browning,
1966; Leonard, 1969). Similarly, resistant plants can act as a barrier to
the transmission of a virus among plants of a susceptible component in a
mixture (Broadbent, 1957; Sandfaer, 1970b). The effects of admixed non-

host plants on insect infestations may however be more subtle. The nonhost
plants may interfere with the visual cues by which the insect finds its
host (International Rice Research Institute, 1974), or with the olfactory
cues for host finding (Tahvanainen and Root, 1972); the nonhost plants
may interfere with feeding behavior (Tahvanainen and Root, 1972), may
attract predators (International Rice Research Institute, 1974), or
provide a more favorable biotic environment for the growth of predator
populations (Muller, in Lampeter, 1960).
If, as in some mixtures of cotton genotypes grown in India (Simmonds,
1962), each of the two components is attacked simultaneously by a disease
to which the other is resistant, theory suggests that pathogen escape in
the mixture may result in the mixture overyielding (Chilvers and Brittain,
1972). If only one component is affected, the mixture is more likely to
yield intermediate between the monocultures. Whether the mixture
yields close to PI (as in Suneson, 1960), or closer to P (as in Klages,
1936) is decided by several factors such as time of onset of the disease,
the degree of pathogen escape due to the dilution effect mentioned above,
and whether the affected component is the aggressor or the subordinate
(de Wit, 1960; Sibma et al., 1964).
Although the above discussion suggests that the growing of appropriate
mixtures will help to minimize the effects of pathogens, it should be pointed
out that mixtures have no automatic advantage in this respect. After a
long series of elegant experiments, Sandfaer (1968, 1970a,b) concluded
that the low RYT values (based on grain yield) in a series of varietal
mixtures of barley were due to barley stripe mosaic virus being transmitted
from a certain variety, a symptomless carrier, to the other, sensitive compo-
nents of the mixtures. The virus caused sterility in the sensitive varieties,
an effect that occurred too late in growth for the carrier variety to show
a compensating increase of grain growth. In this context, an earlier study
of van den Bergh and Elberse (1962) may be mentioned. Here, low values
of dry-matter RYT in a series of grass mixtures were attributed to an ex-
actly similar situation; the evidence for the involvement of the virus was,
however, only circumstantial. In connection with diseases caused by fungi,
Butler and Jones (1949) stated that, in general, more active carbon assimi-
lation (due for instance to increased light levels) increases susceptibility to
obligate parasites of the green parts of plants (e.g., rusts). According to
Gaumann (1950), however, reduced light levels lead to increased suscepti-
bility to “eusymbiotic” parasites (e.g., Fusarium spp.). With this in mind,
it is not difficult to imagine a crop mixture in which, owing to neighbor
effects, each of the components was more susceptible than in monoculture
to a parasitic fungus which was attacking it. However, so far only single
204 B. R. TRENBATH
components have been reported to be worse infected in mixture, e.g., al-

falfa by Rhizoctonia solani when mixed with a grass (Chamblee, 1958)
and rice by blast when mixed with corn (International Rice Research Insti-
tute, 1974).
Before passing to the final conclusions, the points discussed in this sec-
tion may be summarized by means of Table 111. When considering the
implications of an observed deviation of RYT from unity unaccompanied
by transgressive yielding, or of a report of transgressive yielding without
the data needed to calculate an RYT, it should be remembered that over-
yielding implies RYT > 1 and underyielding implies RYT < 1 but that
the converses are true only if the monoculture yields are sufficiently similar.
TABLE 111
Summary of the Evidence Discussed in Section V Concerning Deviations of
Relative Yield Totals (RYT) from Unity and Transgressive Yielding.
RYT >1
without
Mechanism that might lead to RYT >1 overyielding Overyielding
Differing growth rhythms

Differing rooting depths
+ +
+" +"
+b
Nutritional complementation
Enhanced light-use efficiency
Enhanced water-use efficiency
+
Allelopathy
Lodging escape
Pathogen escape
RYT < 1
without
Mechanism that might lead t o R Y T <1 underyielding Underyielding
Lowered light-use efficiency

Allelopathy
Induced lodgingd
Disease expression through infection of a susceptible
component (+)
A cross shows that a t least one report exists t o substantiate the operation of the
given mechanism to produce the given result. I f , for lack of data o n biomass, n report
concerning grain yield has been cited, the cross is shown in parentheses.
Unreplicated, but 5 out of the 6 mixtures grown overyielded.
Only in mixtures of legume + nonlegume.
If lodging in the mixture were to follow the pattern in the more susceptible
monoculture.
VI. Conclusions
The main findings of this review are now listed:

1. Most binary mixtures have been recorded as yielding at a level be-
tween the yields of the components’ monocultures (see Table I ) . This
“nontransgressive” yielding is what might be predicted on the assumption
of competition between components for the same resources. Such competi-
tion would be expected, as a first approximation, to lead to equal propor-
tional increases and decreases of plant biomass compared with per-plant
performance of the components in monocultures. This implies that the rela-
tive yield totals (RYT’s) of mixtures would have values close to unity. This
is found in practice (see Table 11).
2. A minority of binary mixtures has been recorded as yielding trans-
gressively, that is to say outside the range defined by the yields of the com-
ponents grown in monoculture. This suggests that the above proportional
model may not always apply, but the frequent lack of repetition of experi-
ments and the small margins by which the mixture yields transgressed the
range between the monoculture yields usually make it impossible to say
whether a given case of transgressive yielding was due to experimental error
or to a real effect. Since a series of mechanisms can be suggested that
could plausibly lead to mutually beneficial effects between mixture compo-
nents, it seems likely, or at least possible, that some of the observed cases
of overyielding are due to such mechanisms. The frequent absence of con-
clusive experimental evidence in favor of the operation of the hypothetical
mechanisms may be due in part to the lack of sustained investigations of
specific cases of overyielding and in part to their extreme sensitivity to
variations of environmental conditions. Also, few informed attempts have
yet been made to increase the likelihood of overyielding by a conscious
choice of conditions and genotypes. Similar remarks apply to underyielding
by mixtures, although rather fewer mechanisms have been suggested to ac-
count for it.
3. The operation of mechanisms resulting in the stimulation or inhibi-
tion of the growth of mixture components beyond expectations based on
the proportional model give transgressive yields only if the monocultures
are sufficiently similar. A convenient measure of such stimulation or inhibi-
tion, and thus of the mechanisms underlying them, is the relative yield
total. This index takes account of the monoculture yields in such a way
as to detect the operation of these mechanisms even in mixtures which do
not yield transgressively.
Owing to the complexity and unpredictability of many agricultural PCO-
systems, the control over them which man claims he has is often only nomi-
206 B. R. TRENBATH
nal. Given a sound theoretical framework, however, he can hope for fruitful
research into ways of improving this control. This survey of literature at-
tempts to point to areas of ignorance and to suggest means of strengthening
our research so that we may seek to proceed towards more effective exploi-
tation of mixed crops.
ACKNOWLEDGMENT
I am grateful to Professor C. M. Donald and Dr. I. Valentine for their critical

reading and discussion of drafts of the manuscript.
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Biomass Productivity of Mixtures

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BIOMASS PRODUCTIVITY OF MIXTURES

Mixtures of field crops are still extensively grown in primitive agricul-

1967). Judged from this standpoint, the performance of mixtures seems

Apart from a section on underlying theory, this review will consider

II. Comparison of Yields of Mixtures and Monocultures

To facilitate description, a number of terms and symbols are introduced.

Grasses 3 Ahlgren and Aamodt

a “mixture diallel” was used. Experiments conforming to this design con-

of the monocultures; the difference between 60.2% and 39.8% is highly

A fourth instance of “significant” overyielding has been reported which,

111. Theoretical Considerations

Let us assume that a 1 : l mixture of the seeds of two genotypes has

If per-plant yields of i and j in the mixture are the same as in their

However, as many early experiments showed (e.g., Montgomery, 1912;

1961). They result in what may be called either “interference effects” or

of a “compensating” type (Aberg et al., 1943; Donald, 1963). If the plant

Quontity of resource acquired / plant

FIG. 1. Graphical interpretation of de Wit's (1960) model when components of

IV. Types of Interaction Causing Nontransgressive Deviations

0.5 to 0.7 to 0.9 to 1.1 to 1.3 to 1.5 to

Grasses 1 2 Ahlgen and

Data derived from a series of cuts.

each other freely, there is a possibility of competition for water and/or

or to be explained by alternative mechanisms (Muller, 1966, but see Bar-

would have drawn false conclusions. The experimental separation of the

V. Mechanisms Capable of Causing Transgressive Yielding by Mixtures

be more productive than any monospecific population. Gause (1934)

of overyielding mixtures were, respectively, 2 and 1 out of 5. The reality

( a ) that canopy structures were not actually measured in any of these

mixture (Broadbent, 1957; Sandfaer, 1970b). The effects of admixed non-

components have been reported to be worse infected in mixture, e.g., al-

Differing growth rhythms

Lowered light-use efficiency

The main findings of this review are now listed:

I am grateful to Professor C. M. Donald and Dr. I. Valentine for their critical

Brouwer, R. 1966. In “The Growth of Cereals and Grasses” (F. L. Milthorpe

Grodzinsky, A. M. 1965. “Allelopathy in Life of Plants.” Naukova Duma,

Trenbath, B. R. 1970. M.Sc. Thesis, University of Wales.

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