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Fruit production and quality of tomato plants (Solanum

lycopersicum L.) are affected by green compost and
arbuscular mycorrhizal fungi
a b c a
A. Copetta , L. Bardi , E. Bertolone & G. Berta
a
Dipartimento di Scienze dell'Ambiente e della Vita , University of Piemonte Orientale
“Amedeo Avogadro” , via Bellini 25/G, 15100, Alessandria, Italy
b
Ente C.R.A., Istituto Sperimentale per la Nutrizione delle Piante , via Pianezza 115, 10149,
Turin, Italy
c
Dipartimento per la Valorizzazione e la Protezione delle Risorse Agroforestali , University
of Turin, via Leonardo Da Vinci 44 , 10095, Grugliasco, Turin, Italy
Published online: 03 Mar 2011.

To cite this article: A. Copetta , L. Bardi , E. Bertolone & G. Berta (2011) Fruit production and quality of tomato plants
(Solanum lycopersicum L.) are affected by green compost and arbuscular mycorrhizal fungi, Plant Biosystems - An
International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana, 145:1,
106-115, DOI: 10.1080/11263504.2010.539781

To link to this article: http://dx.doi.org/10.1080/11263504.2010.539781

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 Plant Biosystems, Vol. 145, No. 1, March 2011, pp. 106–115

Fruit production and quality of tomato plants (Solanum lycopersicum

L.) are affected by green compost and arbuscular mycorrhizal fungi

A. COPETTA1, L. BARDI2, E. BERTOLONE3, & G. BERTA1

1
Dipartimento di Scienze dell’Ambiente e della Vita, University of Piemonte Orientale ‘‘Amedeo Avogadro’’, via Bellini 25/G,
15100, Alessandria, Italy, 2Ente C.R.A., Istituto Sperimentale per la Nutrizione delle Piante, via Pianezza 115, 10149,
Downloaded by [Ams/Girona*barri Lib] at 06:38 14 November 2014

Turin, Italy, and 3Dipartimento per la Valorizzazione e la Protezione delle Risorse Agroforestali, University of Turin, via
Leonardo Da Vinci 44, 10095, Grugliasco, Turin, Italy

Abstract
Green compost, produced from green wastes, is characterized by high quality standards. The influence of soil amendment
with green compost and with an arbuscular mycorrhizal (AM) fungal-rhizobacterial mixed inoculum on plant growth and
on yield and quality of fruits in tomato was studied. Inoculated and non-inoculated plants were transplanted in pots with
soil added with different compost proportions (0, 25, 50, 75 and 100%). Tomato fruits were analyzed for glucose, fructose,
nitrate, nitrite, malate, citrate, carotenoids, and ascorbic acid content. There are significant differences in shoot and root
dry weights between mycorrhizal and control plants, whereas the addition of compost to substrate until 75% increased
shoot dry weight. AM tomato plants showed significantly higher concentrations of glucose and malate in fruits. Citrate was
higher with 50% and 75% compost, while its lowest concentration was detected in plants without compost. Nitrate content
was increased by AM inoculum and compost; nitrite content was decreased. Higher amounts of compost increased
carotenoid content; ascorbate levels were significantly higher in control plants. These results show that the use of AM
inoculum and green compost can improve fruit quality, affecting biochemical composition and relative proportion of
various compounds.

Keywords: Arbuscular mycorrhizal fungi, carotenoids, fruit quality, green compost, Solanum lycopersicum L., sugars

compounds (carotenes, lutein, and b-cryp-tox-

Introduction
Current methods in fruit and vegetable production
use fertilizer application regimes that are aimed to
reach maximum volume or weight of yield, some-
times neglecting the aspects of quality (Oke et al.
2005).
Tomato (Solanum lycopersicum L.) is one of the
most widely consumed vegetable crops in the world
and a main component of the traditional Mediterra-
nean diet (Dumas et al. 2003). Since tomato plays an
important role in human diet, its nutritional quality
is of particular concern to consumers throughout the
world (Chapagain & Wiesman 2004). Tomato fruits
represent a major source of antioxidant micronu-
trients such as vitamin C, lycopene, and b-carotene.
These compounds have been shown to play a role in
the prevention of diseases: lycopene and other
anthin) in tomatoes reduce prostate cancer risk
(Giovannucci et al. 1995), cardiovascular patholo-
gies (Limpens et al. 2006) and they could be
responsible for the lower risk of developing digestive
tract pathologies (Franceschi et al. 1994). Particu-
larly, lycopene suppresses cell proliferation (Levy
et al. 1995) and interferes with cancer cell growth
(Clinton et al. 1996). The relative contributions of
taste and aroma to tomato flavor has not been clearly
delineated, but sweetness is usually correlated to
total soluble solids and sugar content, mostly
fructose and glucose, whereas acidity is correlated
to pH and organic acid content, mostly citric and
malic acid (Malundo et al. 1995; Auerswald et al.
1999). Tomato fruit taste and antioxidant content
vary among cultivars, growing environments, pro-
duction methods, harvest time, storage, and trans-

Correspondence: Graziella Berta, Dipartimento di Scienze dell’Ambiente e della Vita, University of Piemonte Orientale ‘‘Amedeo Avogadro’’, via Bellini 25/G,
15100, Alessandria, Italy. Tel: þ39-0131-360-232. Fax: þ39-0131-360390. E-mail: graziella.berta@unipmn.it
ISSN 1126-3504 print/ISSN 1724-5575 online ª 2011 Società Botanica Italiana
DOI: 10.1080/11263504.2010.539781

port modalities (Islam et al. 1996; Veit-Köhler et al.
1999; Cano et al. 2003; Dumas et al. 2003; Gautier
et al. 2005; Heeb et al. 2005).
Methods for the production of fruits and vegeta-
bles of optimal quality involve constantly changing
strategies to incorporate the new knowledge coming
from the sciences of production, food quality,
nutritional values, and health beneficial roles of food
components (Oke et al. 2005).
Among the suitable practices for sustainable
management of agriculture soils, there is the use of
arbuscular mycorrhizal fungi (AM fungi). Arbuscular
mycorrhizae are the most common symbiotic asso-
ciation between plants (more than 80%) and soil
fungi (Smith & Read 1997) of the Phylum Glomer-
omycota (Schüßler et al. 2001). Principally, AM
fungi improve mineral nutrition, plant growth,
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tolerance to water stress and adverse environmental
conditions, and resistance to plant pathogens. AM
fungi colonize plant roots, but they influence the
whole plant physiology altering hormonal profile
(Allen et al. 1980; Dixon et al. 1988; Torelli et al.
2000) and influencing photosynthetic efficiency (Rai
et al. 2008). Recent studies have demonstrated that
AM fungi can also affect the volatile compounds
produced in the leaves (Guerrieri et al. 2004) and
increase the concentration of essential oils within
basil (Copetta et al. 2006) and oregano leaves
(Khaosaad et al. 2006).
The use of AM inocula in agriculture is relatively
recent; their efficacy can be influenced by the
modality of application to soil and by the crop. The
success of an AM inoculum can be influenced by the
soil physical structure, porosity, water capacity,
nutrient, and organic matter content (Azcón-Aguilar
& Barea 1997). These soil characteristics are
improved by the addition of compost (Jakobsen
1996; Ros et al. 2003; Pagliai et al. 2004).
Composting has been defined as aerobiotic microbial
activity leading to decomposition of most biodegrad-
able materials, usually mixtures of organic materials,
which results in organic residue stability (Valdrighi
et al. 1996). The addition of compost to soil is
considered an environmentally safe and agronomi-
cally advantageous process at acceptable operational
costs (Dell’Abate et al. 2000). Green compost has a
positive effect on soil productivity and soil character-
istics (Garcı́a et al. 1991) and its use is suitable
mainly for its low risk of toxicity (Witter & Lopez
Real 1998).
The AM-induced improvement of quality and
quantity of plant production can also be emphasized
by the association with rhizobacteria. Among the
free-living microorganisms, plant growth-promoting
rhizobacteria (PGPR) have received special atten-
tion. PGPR can exert a beneficial effect on plant
growth by suppressing soil-borne pathogens (Weller
Compost and AMF affect tomato quality 107
1988), improving mineral nutrition (Kapulnik 1991;
Gamalero et al. 2004), and phytohormone synthesis
(Glick 1995; Gamalero et al. 2008).
The aim of this work was to evaluate the influence
of an AM fungal-rhizobacterial mixed inoculum
associated to green compost on plant growth on
yield quality of fruits of tomato grown in greenhouse;
we analyzed the effects on plant growth and
morphology, and content of sugars, organic acids,
caroteinoids, ascorbic acid, nitrates and nitrites in
fruits.
Materials and methods
Microbial inoculum
Mycorrhizal inoculum consisted of sporocarps,
spores, hyphae, and root fragments colonized by a
number of AM fungi (Glomus mosseae, G. caledonium,
G. viscosum, G. intraradices and G. coronatum),
combined with attinomycetes and rhizobacteria.
Soil and compost
The soil was obtained from a field which was
cultivated with maize, sited in Castelceriolo (Ales-
sandria, Italy). The green compost was provided by
Saceccav (Castelceriolo, Alessandria, Italy) and
consisted of garden and growing refuse composting.
Table I shows physico-chemical characteristics of soil
and green compost.
Seedling production
Tomato seeds (S. lycopersicum L. cv. Guadalete),
kindly provided by Dr. Scaravonati (Solanum c.s.a,
Lodi, Italy), were surface sterilized by gently
shaking in a 5% NaClO solution for 3 min and
rinsed six times for 5 min and four times for
20 min in sterile demineralized water (Gamalero
et al. 2004). The seeds were pregerminated on
moist sterile filter paper at 248C for 3 days and
transplanted into alveolar boxes (50 cm 6 60 cm, 5
cm 6 5 cm; 120 alveoli; 20 cm3 of volume each)
with sterile substrate constituting to 1:1 fine quartz
sand (0.6–1.2 mm):vermiculite (Punto Elle, Turin,
Italy) or 1:1:2 fine quartz sand:vermiculite:mycor-
rhizal inoculum. Plantlets were kept in a growth
chamber with a 16/18 h light/dark photoperiod, 26/
228C light/dark thermoperiod, 150 mEm72s72 light
irradiance at container height, and watered to soil
saturation with tap water. Five plantlets per
treatment were collected 21 days after sowing
and used for morphological analysis; 50 plantlets
were collected after 28 days of growth to
be transplanted in pots for the greenhouse
experiments.
 108 A. Copetta et al.

Table I. Physico-chemical characteristics of soil and green Table II. Treatments.

compost.
Field soil : green compost Non-inoculated Inoculated
Field Green
Parameters soil compost 1 : 0 C M
3 : 1 25% 25%M
pH 6.21 8.02 1 : 1 50% 50%M
Specific Oxygen Uptake nr 0.33 1 : 3 75% 75%M
Rate (mgOgSV h) 0 : 1 100% 100%M
Texture (g/kg)
Sand nr 833
Large silt nr 56
Small silt nr 69 Plant morphology and growth analysis
Clay (g/kg) nr 42
Dry residue (1108C) 990 402.2 The following parameters were determined on 21
Moisture content (g/kg) 10 597.8 days old plantlets: shoot length, leaf number, root
Ashes (6508C) (ss) 977.6 738.1 and shoot fresh weight, total root length, mycorrhizal
Organic substance (ss) 22.4 261.9 colonization and arbuscular density. Roots and
Total elemental content
shoots of 5 months-old plants were dried at 608C
C (g/kg ss) nr 314.2
for 3 days, then weighed to determine the dry weight.
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N (g/kg ss) 50.1 25.9

P (P2O5) (g/kg ss) 0.026 8.7
K (g/kg ss) 0.237 nr
Cd (mg/kg) nr 0.050 + 0.004 Mycorrhizal colonization
Cr (mg/kg) nr 0.019 + 0.002 Forty randomly chosen 1 cm-long pieces were cut
Ni (mg/kg) nr 0.200 + 0.021
Pb (mg/kg) nr 6.338 + 0.279 from each root system and fixed in 70% ethanol,
Toxicity test then stored at 48C until mycorrhizal colonization
Heterocypris incongruens nr Biostimulation analysis.
Dictyostelium discoideum nr Feebly toxic Mycorrhizal colonization and arbuscule abun-
Vibrio fischeri nr Feebly toxic dance were estimated according to Trouvelot et al.
Eisenia parva nr No toxic
Lepidium sativum nr Biostimulation
(1986). Root pieces were cleared for 30 min at 608C
Sorghum vulgare nr No toxic in 10% KOH, stained with 1% methyl blue in lactic
Cucumis sativus nr Biostimulation acid and mounted on a slide. The intensity of
Allium porrum nr No toxic mycorrhizal colonization was evaluated microscopi-
Zea mays nr No toxic
cally.
nr, not recording.
Fruit analysis
Fruits of same dimension (approximately 20 g) were
Greenhouse assay experimental design and growth
harvested at the red ripe stage and stored at –208C.
condition
Frozen fruits were deprived of skins and homoge-
Twenty-eight days-old inoculated and non-inocu- nized with mortar and pestle. Homogenized fruit
lated tomato seedlings were transplanted in pots with tissue was centrifuged (10,000 rpm 6 5 min) and the
soil added with different compost proportions and following parameters were determined in the super-
cultivated for 5 months (May–September 2004) in a natant: pH by means of a pH meter (Crison
greenhouse sited in Castelceriolo (Alessandria, MicropH 2001, Carpi, Modena Italy) and EC by
Italy). means of an EC meter (Orion Model 120). Twenty-
The experimental design was a block of 50 plastic five grams of homogenized samples were dried at
pots of 7.5 l. Green compost was diluted with field 608C for 3 days. Water and dry matter content were
soil to create compost concentrations of 0, 25, 50, estimated by the difference in the weight before and
75, and 100% (v/v). Ten replicates were made for after lyophilization.
each dilution. Each pot was planted with one Glucose, fructose, nitrate, nitrite, malic, citric, and
inoculated or non-inoculated plantlet (five contain- ascorbic acid content were determined using stan-
ers per treatment – see Table II). During transplant- dard enzimatic analytical kits (R-Biopharm, Roche)
ing mycorrhizal (M), plant roots were surrounded for the detection of these compounds in foods. One
with 1 l of 1:1 field soil:mycorrhizal inoculum gram of tomato juice was diluted to 100 ml with
mixture in order to reinforce the inoculum. Plants MilliQ water (Millipore). The suspensions were
were shaded from May to September and watered filtrated through Whatman filter paper and the clean,
every day for 8 min at 4 a.m. with running tap water. colorless serum was used for analysis.
Plants were harvested after 5 months from the The analysis of carotenoids was carried out
sowing. according to Oke et al. (2005) for the extraction in
 Compost and AMF affect tomato quality 109

organic phase and to Hart and Scott (1995) for the

Mycorrhizal colonization
chromatographic analysis. Briefly, 4 g of tomato
pulp, after mortar crushing, were put and weighed Mycorrhizal colonization and arbuscule formation
into 250 ml brown bottles. In each bottle of significantly decreased with the increase of green
100 ml 2:1:1 hexane: acetone: ethanol solution
containing 0.1% butylated hydroxytoluene (BHT) Table III. Morphological parameters of control and AM plant after
and 1 mg/ml of b-apo-80 -carotenal dissolved in 21 days of growth.
hexane as internal standard were added; the bottles
were closed and shaked for 10 min at 250 rpm on Parameters Control AM plant
an orbital shaker. After adding 15 ml of distillated Leaf number 4.0 + 0.1 a 5.2 + 0.4 b
water and a further 5 min shaking, the organic Shoot length (cm) 4.1 + 0.3 a 6.4 + 0.6 b
layer (50 ml) was separated from the aqueous Shoot fresh weight (g) 0.112 + 0.007 a 0.164 + 0.012 b
layer (65 ml) and centrifuged at 4000 rpm for Root fresh weight (g) 0.062 + 0.003 a 0.082 + 0.008 b
Mycorrhizal nd* 15.3 + 2.1 b
6 min.
colonization (%)
The organic extract was directly subjected to Arbuscule nd 13.4 + 1.5 b
HPLC analysis (Jasco PU-980 equipped with a Jasco abundance (%)
UV-2075 Plus detector) on a Supelco LC-18
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column, 150 mm 6 4.6 mm, particle size 5 mm. *nd, not detected. Inoculum application increased leaf, shoot and
root parameters. No sign of AM colonization was detected in
The operating condition were: isocratic elution with control roots. Mean values + SEM are shown; different letters
mobile phase acetonitrile:methanol:dichloromethane indicate statistically significant differences (p 5 0.05) between
75:20:5 containing 0.1% BHT and 0.05% trietha- control and mycorrhizal plants, along the lines of the table.
nolammine; methanol was added with 0.05 M
ammonium acetate; flow rate 1.5 ml/min, injection Table IV. Shoot and root dry weights (g) of tomato plants after five
volume 20 ml, 450 nm. Lycopene and b-carotene months of growth.
were quantified by external standards analysis. All
Shoot dry weight Root dry weight
reagents were purchased from Sigma-Aldrich Cor-
poration (USA). C 4.82 + 0.49 a 1.01 + 0.13 a
25% 22.84 + 2.23 bc 6.63 + 1.16 bc
50% 27.08 + 0.71 b 7.44 + 1.21 bc
Statistical analysis 75% 26.72 + 4.72 b 6.51 + 1.47 bc
100% 29.68 + 3.29 bd 6.07 + 1.03 bc
Data were statistically analyzed by ANOVA followed M 17.28 + 2.25 ce 4.38 + 0.88 cd
by Fisher’s probable least-squares difference test 25% M 26.82 + 3.55 bf 5.54 + 0.89 bcd
with cut-off significance at p ¼ 0.05. 50% M 24.45 + 6.16 be 3.44 + 0.95 ad
75% M 36.96 + 4.09 d 5.74 + 0.82 bcd
100% M 35.32 + 1.70 df 5.86 + 1.16 bcd
Results Mean values + SEM are shown; different letters indicate statisti-
cally significant differences (p 5 0.05) among treatments, along
Seedling growth
the columns of the table.
After 21 days of growth, plants treated with microbial
inoculum were colonized, whereas no AM coloniza-
tion was detected in control roots. Mycorrhizal
Table V. AM colonization parameters of tomato plants.
inoculum induced significant increases in leaf
number, in shoot length and shoot and root fresh Mycorrhizal Arbuscule
weight (Table III). degree % abundance %

C nd nd
Plant growth 25% nd nd
50% nd nd
After 5 months of growth, there were a number of 75% nd nd
differences between plants amended with mycorrhi- 100% nd nd
M 37.0 + 4.4 b 24.7 + 5.1 b
zal inoculum and/or green compost and control
25% M 22.6 + 10.0 bc 17.2 + 5.6 bc
plants (Table IV). There were significant differences 50% M 13.5 + 8.6 c 6.7 + 3.6 c
in shoot and root dry weights between mycorrhizal 75% M 13.2 + 9.4 c 8.8 + 5.3 c
(M) and control (C) plants. Shoot biomass signifi- 100% M 0.5 + 0.5 a 0.2 + 0.2 a
cantly increased with the higher green compost
nd, not detected. No sign of infection was detected in control
content in the growth substrate. The best dry weight roots. Mean values + SEM are shown; different letters indicate
yield occurred at compost rates of 75% and AM statistically significant differences (p50.05) among treatments,
fungi application. along the columns of the table.
 110 A. Copetta et al.

Table VI. Nutrient content (g/kg) of tomato in several treatments.

Glucose Fructose Malate Citrate Ascorbate* Nitrite Nitrate

C 3.0 + 0.3 a 4.3 + 0.3 a 0.7 + 0.1 a 2.2 + 0.1 ad 799 + 138 a 1.91 + 0.81 ab 2.43 + 0.86 a
25% 2.5 + 0.4 a 5.5 + 1.2 ab 0.7 + 0.1 a 3.1 + 0.4 ac 457 + 96 b 2.23 + 0.49 a 0.83 + 0.75 a
50% 3.7 + 1.0 a 3.9 + 0.7 a 1.0 + 0.1 c 4.4 + 0.6 b 126 + 50 cd 1.43 + 0.81 ab 3.20 + 1.46 a
75% 2.7 + 0.6 a 4.8 + 0.5 ab 1.1 + 0.2 c 3.2 + 0.3 c 81 + 24 cd nd 10.91 + 2.88 b
100% 2.2 + 0.3 a 3.4 + 0.6 a 1.0 + 0.2 ac 2.8 + 0.4 acd 113 + 66 cd 0.07 + 0.07 c 14.05 + 3.75 b
M 5.9 + 1.0 b 6.4 + 0.6 b 1.8 + 0.3 b 1.9 + 0.2 d 250 + 84 bd 0.73 + 0.40 bc 13.10 + 3.23 b
25% M 2.2 + 0.2 a 3.9 + 1.0 a 1.1 + 0.3 ac 2.3 + 0.2 ad 21 + 20 c nd 11.25 + 0.61 b
50% M 3.3 + 0.8 a 4.3 + 0.6 a 1.5 + 0.3 bc 2.3 + 0.3 ad 56 + 22 cd nd 12.06 + 0.52 b
75% M 2.9 + 1.0 a 3.4 + 0.7 a 1.1 + 0.1 ac 2.4 + 0.3 acd 162 + 115 cd 0.23 + 0.12 c 13.15 + 1.13 b
100% M 2.4 + 0.1 a 3.5 + 1.2 a 1.5 + 0.3 bc 2.6 + 0.2 ac 103 + 32 cd 0.11 + 0.11 c 14.87 + 0.22 b

*expressed in mg/l. nd, not detected. Mean values + SEM are shown; different letters indicate statistically significant differences (p 5 0.05)
among treatments, along the columns of the table.

Table VII. pH, water content and conductance in juice of tomato ences among the treatments in tomato juices pH.
fruits. Water content was significantly lower in fruits of
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Water Conductance
plants grown in pots with 25% and 50% green
pH content (g/kg) (mS) compost than in fruits of control plants. Analysis of
conductance of tomato juice showed a significantly
C 3.95 + 0.03 a 934 + 5 a 5.1 + 0.2 a higher concentration of minerals in plants treated
25% 3.99 + 0.02 a 913 + 4 b 4.7 + 0.5 a
with AM inoculum and/or green compost than in
50% 4.02 + 0.02 a 914 + 7 b 6.9 + 0.6 b
75% 4.00 + 0.02 a 918 + 14 ab 7.0 + 0.4 bd control plants (Table VII).
100% 3.97 + 0.02 a 927 + 5 ab 7.2 + 0.3 bd Lycopene was the most abundant carotenoid of all
M 3.96 + 0.02 a 927 + 5 ab 6.0 + 0.1 c samples of tomato fruits, followed by b-carotene. A
25% M 3.99 + 0.03 a 928 + 4 ab 5.9 + 0.2 c higher content of green compost in plant growth
50% M 4.08 + 0.10 a 935 + 12 a 6.7 + 0.4 bcd
substrate increased b-carotene, lycopene, and luteine
75% M 4.00 + 0.03 a 935 + 6 a 7.1 + 0.6 cd
100% M 3.99 + 0.01 a 936 + 5 a 7.7 + 0.4 d content of fruits (Table VIII).

Mean values + SEM are shown; different letters indicate statisti-

cally significant differences (p 5 0.05) among treatments, along
the columns of the table.
compost in the soil (Table V). No AM colonization
was detected in uninoculated roots.
Fruit quality
In tomato fruits fructose and citrate content was
always higher than glucose and malate content,
respectively. Several differences in nutrient content
were detected among the treatments.
Fruits of inoculated plants (M) contained a
significantly larger quantity of glucose than fruits of
all other treatments (Table VI). In this treatment
(M), tomatoes had also the largest fructose content.
Malate content of fruits significantly increased with
inoculum application and with green compost
content higher than 50%. Tomato fruits of plants
grown with 50% green compost had significantly
larger citrate content than the other treatments.
Application of green compost and/or microbial
inoculum in the substrate significantly decreased
ascorbate content in fruits. Moreover, microbial
inoculum and green compost decreased and in-
creased respectively the nitrite and the nitrate
content of fruits. There were no significant differ-
Discussion
The growth effect in inoculated plantlets, compared
to control ones, was already reported in other studies
on tomato. Berta et al. (2002) showed that G.
mosseae BEG12 improves root development improv-
ing input and transport of nutrient. Co-inoculation
of G. mosseae and rhizobacteria synergistically in-
creases shoot and root growth, and influences root
architecture, improving phosphate nutrition (Gama-
lero et al. 2004).
After 5 months of growth, tomato plant growth
was correlated to compost content in the substrate.
Recent researches (Valdrighi et al. 1996; Atiyeh et al.
2000; Ouédraogo et al. 2001; Cooperband et al.
2002) about several horticultural and economical
species growing in substrates with different soil/
compost ratio confirm the beneficial effects of
compost on shoot and root development. A sig-
nificant increase of several parameters, e.g. length
and height of shoot, leaf number and length of
inflorescence, in Solanum melongena, Cyamopsis
tetragonoloba, Capsicum annuum and S. lycopersicum
harvested after growth in soil/compost substrate was
observed (Gajalakshmi & Abbasi 2002).
At harvest time, inoculated tomato roots were
mycorrhized, but the increase of green compost in
the culture substrate decreased root colonization

Table VIII. Tomato carotenoids in fruits of control and AM plants.
b-carotene (mg/kg)
C 57.2 + 4.5 a
25% 59.8 + 16.3 a
50% 83.0 + 10.8 acd
75% 109.5 + 16.9 bcd
100% 147.8 + 8.5 b
M 56.3 + 6.9 a
25% M 90.1 + 22.4 ad
50% M 107.1 + 12.9 bd
75% M 108.0 + 29.4 bd
100% M 145.1 + 13.9 b
Green compost improved carotenoids content. Mean values + SEM are shown; different letters indicate statistically significant differences
(p50.05) among treatments, along the columns of the table.
degree. The effects of compost on AM fungi are
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contrasting. Some compost types (from selected
waste) improve the growth of this simbionts.
Particularly, 20% of vermicompost in the soil
improve sorghum and strawberry growth and AM
colonization (Murphy et al. 2000; Cavender et al.
2003). Gryndler et al. (2005) show that humic
substances into green amendment stimulate the
development of AM hyphae. Other organic amend-
ments do not influence the formation of AM
structure (Leporini et al. 1992; Hafner et al. 1993;
Wani & Lee 1995). Compost can inhibit AM fungi
infection: compost increase negatively affects G.
mosseae and G. deserticola colonization in soy and
lettuce roots (Martı́n et al. 2002).
Several factors can have influenced the low
mycorrhization observed in tomato roots growing
in pots with green compost concentration higher
than 25%. The amendment use enhanced phosphor-
ous (P) content in soil culture (Toor & Bahl 1997).
Several studies show a negative effect of the increase
of available P on AM populations; moreover the
suppressive effect of P on AM colonization (Allen
et al. 1980; Valentine et al. 2001; Fusconi et al.
2005) and spore density (Menge et al. 1978) is
widely documented. Duke et al. (1994) observed a
reduction of arbuscular formation in zones of
Artemisia tridentata roots exposed to high P concen-
trations. Experiments carried out to develop pro-
cesses for the production of AM fungus inocula,
show that 1:4 and 1:9 ratios of yard clippings
compost and vermiculite tend to yield the greatest
development of colonization of bahiagrass roots
(Douds et al. 2006), but the P content of compost
used in that assay was smaller than that of our study.
Moreover, composted organic material contains
products of decomposition, which can have strong
inhibitory effects on mycorrhizal fungi (Gryndler
et al. 2008).
Also the presence of PGPR (e.g. Attinomyces,
Pseudomonas spp., Bacillus spp.) in mycorrhizal
Compost and AMF affect tomato quality 111
Lycopene (mg/kg) Luteine (mV)
449.9 + 81.0 a 21.9 + 2.2 a
455.8 + 62.3 a 25.2 + 4.5 a
701.9 + 207.8 ab 46.9 + 11.2 bcd
607.5 + 151.2 ab 38.3 + 5.8 ac
975.8 + 122.1 b 61.2 + 6.5 bc
504.7 + 81.2 ac 30.6 + 3.9 ad
530.7 + 89.0 ac 36.1 + 4.6 ade
760.0 + 225.9 ab 35.6 + 5.3 ade
876.5 + 196.5 bc 49.2 + 7.3 bce
839.1 + 105.6 bc 55.0 + 5.1 b
inoculum can synergically improve tomato growth
(Gamalero et al. 2004). Poulton et al. (2001, 2002)
showed that mycorrhizal tomatoes produce more
seeds and fruits compared to control ones; moreover
the symbiosis improved leaf area, flower number,
female and male reproductive functions, fruit num-
ber, and seed quality. These results show that AM
infection can increase the fitness of host species
influencing the reproductive functions. Meunchang
et al. (2006), adding compost in the substrate,
observed an increase in tomato shoot height, and a
further increase adding the rhizobacteria Azospir-
illum, Azotobacter and Beijerinckia.
Many compounds (sugars, acids and more than
two hundred volatile constituents) determine the
flavor and fragrance of tomato fruits, but it is not yet
fully clear how and to what extent they contribute to
flavor and aroma (Malundo et al. 1995; Auerswald
et al. 1999). Moreover, growing environmental
factors and many other factors influence tomato
fruit taste.
Sweetness is particularly appreciated in tomatoes
for industrial use, and is usually correlated with
fructose and glucose content mainly accumulated
into the vacuole of fruit cells (Ordóňez et al. 2005).
Our results show that the fructose content of fruits is
higher than the glucose one. Other studies confirm
these data (Baldwin et al. 1991; Islam et al. 1996).
Malundo et al. (1995) showed that an increase of
fruit sugars linearly improved tomato flavor for a
sample of consumers. As shown in Table VI, we
observed a strong increase of both glucose and
fructose in fruits from inoculated plants in absence of
compost (M) in respect to all the other samples; so
we can deduce that microbial inoculum significantly
improved the tomato flavor. The taste experience is
subjective and often is not the single factor used to
determine the fruit quality (Cardello 1995; Shewfelt
1999). Generally nutritional, microbiological, and
physico-chemical characters determine food quality.
Moreover, food quality is a consumer-based
 112 A. Copetta et al.
perceptual/evaluative construct that depends on
person, place and time and is subjected to the same
influences of context and expectations as are other
perceptual / evaluative phenomena (Cardello 1995;
Moskowitz 1995; Shewfelt 1999). It is important to
emphasize that there are many difficulties to estimate
the importance of several compounds in tomato
flavor and fragrance.
Acidity is correlated with pH and acid content,
and citrate concentration is usually higher than
malate in tomato (Stevens 1979; Malundo et al.
1995; Auerswald et al. 1999). Our data confirm these
observations. Moreover inoculated plants (M) pro-
duced fruits with malate content significantly higher
compared to nearly all treatments. Citrate content
significantly increased in sample from plants growing
in soil:green compost 1:1, and it decreased when
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inoculum was added into substrate.
Ascorbate, as vitamin E, flavonoids, anthocians,
phenolic compounds and carotenoids, is an impor-
tant antioxidant. In our work, ascorbic acid (vitamin
C) concentration was affected by both increasing
green compost and microbial inoculum. Conditions
and plant nutrition can influence vitamin C content
in fruits. For example, rising N fertilization decreases
ascorbic acid concentration in lemon, tomato, and
potato (authors cited in Veit-Köhler et al. 1999).
Moreover, mycorrhizal fungi can improve N nutri-
tion in host plants. Inorganic N is quickly incorpo-
rated into aminoacids and organic compounds and
translocated into the roots (Barea et al. 1987; Frey &
Schüepp 1993; Tobar et al. 1994; Ibijbijen et al.
1996; Azcón & Tobar 1998; Faure et al. 1998). In
our work, the increase of green compost/soil ratios in
pots corresponded a nitrite and nitrate content
decrease and increase, respectively, in tomato fruits;
while fruits of control plants contained lower
concentrations of inorganic nitrogen and higher
quantities of ascorbate (Table VI). Mycorrhizal
plants, absorbing reduced nitrogen, can conserve
energy useful to synthesize secondary metabolites
related to flavor and quality of fruits as previously
hypothesized by Heeb et al. (2005) who observed an
improvement in sweetness, acidity and aromaticity in
fruits of plants fertilized with NH4þ compared to
ones treated with NO37.
Our results show that mycorrhizal inoculum did
not improve carotenoid content of fruits while the
green compost increased b-carotene, lycopene, and
luteine. Chromatograms to carotenoid acknowledg-
ment and analysis are similar to other studies (Hart &
Scott 1995; Lewinsohn et al. 2001), while b-carotene
and lycopene concentration is similar to that found
in well ripe tomatoes by Giuntini et al. (2005). In a
recent study, Mena-Violante et al. (2006) observed a
significant increase in carotenes and xanthophylls of
C. annuum fruits treated with a AM fungi consortium
compared to noninoculated control, while a different
mixture of mycorrhizal fungi and Gl. fasciculatum
treatment did not affect pigment content in chile
ancho fruits. These results show that AM fungi
isolates or mixtures may increase carotenoid content
in fruits. Carotenoid accumulation in plant tissue
appears to be shaped by the physiological, genetic,
and biochemical attributes of a plant species, as well
as by environmental growth factors such as light,
temperature, and fertility (Kopsell & Kopsell 2006).
Some aroma compounds, namely, the apocarotenals
are derived from the breakdown of carotenoids
mainly lycopene and b-carotene (Lewinson et al.
2005 a,b). Moreover, correlations between lycopene
and the content of aroma volatiles have been
reported in cherry tomatoes (Ishida et al. 1998).
Changes in nutritional fertility affect plant carotenoid
accumulations. Nitrogen fertilization seems to im-
prove carotene accumulation. In fact, increasing N in
the nutrient solution increased lutein–zeaxanthin, b–
carotene content in parsley (Petroselinum crispum
Nym.) (Chenard et al. 2005). Hochmuth et al.
(1999) reported that carrot root carotenoid concen-
tration is maximized with 160 kg ha–1 N. Moreover
those N fertilization rates also maximized carrot root
yield and sugar content. b-Carotene, lycopene and
luteine have been shown to play a role in the
prevention of human diseases. b-Carotene is biolo-
gically separated to produce vitamin A, required for
vision, epithelia maintenance, secretion of mucus,
and reproduction (Mayne 1996). Lycopene com-
bined with vitamin E may inhibits the growth of
prostate cancer (Limpens et al. 2006). Lutein is a
carotenoid selectively deposited in the human retina
and lens, and responsible for the yellow pigmentation
(macular pigment) (Bone et al. 1997; Khachik et al.
1997). This pigment is postulated to participate in
photoprotection, and diminished macular pigment
levels may be related to retinal damage (Mares-
Perlman & Klein 1999; Wooten et al. 1999). The
consistently lower risk of cancer for a variety of
anatomic sites that is associated with higher con-
sumption of tomatoes and tomato-based products
adds further support for current dietary recommen-
dations to increase fruit and vegetable consumption
(Giovannucci 1999).
This work shows that both mycorrhizal inoculum
and green compost improved growth and yield of
tomato plant, did not modify the ascorbic acid
content of fruits, decreased the nitrite and increased
the nitrate concentration (nitrate and nitrite contents
of fruits were always lower than legal limits – data not
shown). Moreover, AM fungi increased the glucose
and malic acid content of fruits whereas green
compost increased fruit carotenoids content. The
application of mycorrhizal fungi and green compost
improving nutrient absorption improved flavor,

nutritional value, and human disease protection of
tomato fruits. In conclusion, AM fungi and the green
compost have the potential to be a useful agricultural
tool for plant improvement and product quality in
biosustainable agroecosystem.
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