265

Adv. Hort. Sci., 2022 36(4): 265­273                                                                                                    DOI: 10.36253/ahsc­13352

Arbuscular mycorrhizal fungi potentiate 
the root system and the quality of  

goldenberry fruits 
 
 
J.L.T. Chiomento 1 (*), D. Filippi 1, G.M. Krasnievicz 2, J.E.C. De Paula 2, M. 
Fornari 3, T.S. Trentin 2 
¹     Postgraduate Program in Agronomy, University of Passo Fundo, Passo 

Fundo, RS, Brazil. 
²    Undergraduate Program in Agronomy, University of Passo Fundo, 

Passo Fundo, RS, Brazil. 
3       Postgraduate Program in Agricultural Engineering, Federal University 

of Santa Maria, Santa Maria, RS, Brazil. 
 
 
Key words: Berry flavor, mycorrhization, Physalis peruviana L., root morphology. 
 
 
Abstract: The lack of information on the horticultural performance of golden­
berry (Physalis peruviana L.) is one of the factors that limits the expansion of 
the crop. Still, aiming to establish a sustainable management for this culture, 
inoculation with arbuscular mycorrhizal fungi (AMF) can be adopted. Therefore, 
the objective of the research was to investigate whether goldenberry plants in 
the absence and presence of inoculation with AMF differ in terms of horticul­
tural performance. The four treatments studied were the absence (control) and 
the presence of three inoculants based on AMF (mycorrhizal community, 
Glomus intraradices and Rhizophagus clarus), arranged in a randomized block 
design, with five replications. Goldenberry plants produced in substrate 
enriched with AMF had a more voluminous root system and a greater amount 
of fine roots. Additionally, the fruits were sweeter and more flavorful when 
produced by plants inoculated with the mycorrhizal community and with R. 
clarus. It is concluded that mycorrhization has no effect on fruit production. 
However, goldenberry plants submitted to mycorrhizal biotechnology enhance 
the chemical quality of fruits and present a more profuse root system. G. 
intraradices is most effective in colonizing the roots of the plant host. 
 
 
1. Introduction 
 
     Goldenberry (Physalis peruviana L., Solanaceae) is a horticultural crop 
native to the Andean highlands that has attracted worldwide attention 
due to its bioactive compounds such as carotenoids, physalins and 
polyphenols (Ramadan, 2011). In addition to promoting the health of con­
sumers, these biomolecules present in fruit extracts have antifungal 
action against phytopathogens, such as Botrytis cinerea Pers., and there­
fore can be widely used in agriculture as a bioinput (Filippi et al., 2020). 
Still, goldenberry stands out for its potential for intensive cultivation 

(*)
 
Corresponding author:  

jose­trevizan@hotmail.com 
 
 
Citation: 
CHIOMENTO J.L.T., FILIPPI D., KRASNIEVICZ G.M., 
DE PAULA J.E.C., FORNARI M., TRENTIN T.S., 2022 
­ Arbuscular mycorrhizal fungi potentiate the root 
system and the quality of goldenberry fruits. ­ 
Adv. Hort. Sci., 36(4): 265­273. 
 
 
Copyright: 
© 2022 Chiomento J.L.T., Filippi D., Krasnievicz 
G.M., De Paula J.E.C., Fornari M., Trentin T.S. 
This is an open access, peer reviewed article 
published by Firenze University Press 
(http://www.fupress.net/ index.php/ahs/) and 
distributed under the terms of the Creative 
Commons Attribution License, which permits 
unrestricted use, distribution, and reproduction 
in any medium, provided the original author and 
source are credited. 
 
 
Data Availability Statement: 
All relevant data are within the paper and its 
Supporting Information files. 
 
 
Competing Interests:  
The authors declare no competing interests. 
 
 
Received for publication 6 July 2022 
Accepted for publication 6 October 2022

AHS 
Advances in Horticultural Science



Adv. Hort. Sci., 2022 36(4): 265­273

266

(Etzbach et al., 2018). A single plant can produce 300 
fruits and the productivity of this horticultural crop 
can reach from 20 to 33 tons per hectare (Yildiz et al., 
2015). Despite the traditional establishment of crops 
in the open field (Muniz et al., 2014), the goldenberry 
cultivation in greenhouse is increasing (Aguilar­
Carpio et al., 2018). This is because greenhouse culti­
vation can help to avoid inconveniences such as 
pests, diseases, rain, strong winds, hail and frost 
(Costa et al., 2016). 
     Similar to the traditional cultivation of other 
Solanaceae, such as tomato and pepper, in order to 
obtain an optimal productive yield of goldenberry, 
producers need to use a large amount of chemical 
inputs, which can contaminate the agroecosystem of 
cultivation (Chiomento et al., 2020 a). There is no 
doubt, therefore, that the establishment of agroeco­
logical agriculture is an important tool for sustainable 
food production, with environmental and socioeco­
nomic benefits (Llano et al., 2018). Thus, an alterna­
tive to minimize the inconveniences in the cultivation 
of goldenberry and start the establishment of sus­
tainable management in this horticultural culture 
corresponds to the use of inoculants based on arbus­
cular mycorrhizal fungi (AMF). As there is a limitation 
regarding the availability of commercial AMF­based 
inoculants available in Brazil (Trentin et al., 2022), 
this reduces the use of mycorrhizal biotechnology in 
the production of goldenberry due to the lack of 
knowledge of this bioinput by the producers. The lim­
itation regarding the availability of commercial inocu­
lants is mainly due to the high cost linked to the pro­
duction technology of this bioinput. 
     AMF (phylum Glomeromycota), a ubiquitous 
group of soil microorganisms, establish symbiotic 
associations with more than 70% of vascular plants 
(Brundrett and Tedersoo, 2018). The literature 
reporting the association between mycorrhiza and 
goldenberry is scarce. For example, under saline con­
ditions, AMF increased fruit growth rate (Miranda et 
al., 2011) and improved berry unsaturated fatty acid 
concentration in response to heavy metal stress 
(Hristozkova et al., 2017). Under water stress, arbus­
cular mycorrhiza increased root dry matter accumu­
lation and improved attributes related to plant gas 
exchange (Reyes et al., 2019). In non­stressful envi­
ronments, it was found that goldenberry plants sub­
jected to mycorrhizal biotechnology produced less 
acidic and tastier fruits (Chiomento et al., 2020 a). 
This scarcity of information demands more research 
to fill the existing gaps regarding the morpho­horti­

cultural performance of goldenberry and their inter­
active effects with mycorrhizas. 
     In Brazil there is only one commercial AMF inocu­
lant available to farmers. The commercial scale pro­
duction of this bioinput has high costs linked to the 
inoculum production technology, such as the estab­
lishment of cultures of AMF species and transport, 
handling and development of the carrier substrate 
(Schlemper and Stürmer, 2014). To avoid some of 
these costs, on‐farm production of inoculants is used, 
with indigenous or exotic AMF isolates, in which the 
technology can be easily transferred to farmers 
(Douds Junior et al., 2012). The process of obtaining 
the on‐farm inoculant can be started using AMF 
infective propagules, such as spores, hyphae and 
parts of colonized roots (Douds Junior et al., 2010). 
     Therefore, based on the hypothesis that mycor­
rhizal biotechnology enhances plant host growth and 
improves fruit chemical quality, here we investigate 
whether the horticultural performance of goldenber­
ry is influenced by the use of AMF­based on‐farm 
inoculants. 
 
 
2. Materials and Methods 
 
Plant material 
     The research was carried out in Passo Fundo (28° 
15’ 46” S, 52° 24’ 24” W), Rio Grande do Sul (RS), 
Brazil, in greenhouses, from August (winter) 2018 to 
July (winter) 2019 . 
     A commercial tray with goldenberry fruits at mat­
uration stage 5 was purchased (ICONTEC, 1998). In 
August 2018, seeds from three randomly chosen 
fruits were selected, transferred to paper towels and 
kept at room temperature until dry. Subsequently, 
these seeds were germinated in plastic gerbox boxes 
containing blotting paper and 0.1 molar (M) potassi­
um nitrate (KNO3) solution. The boxes were stored in 
a  b i o c h e m i c a l  o x y g e n  d e m a n d  ( B O D )  o v e n ,  a t 
25°C±1°C, until the plants were obtained for the pro­
duction of seedlings, which constituted the plant 
material for the research. The steps for obtaining the 
plants are shown in figure 1. 
 
Experimental design 
     The four treatments studied were the absence 
(control) and the presence of three inoculants based 
o n  A M F  [ m y c o r r h i z a l  c o m m u n i t y ,  G l o m u s 
i n t r a r a d i c e s  N . C .  S c h e n c k  &  G . S .  M r .  a n d 
Rhizophagus clarus (T.H. Nicolson & N.C. Schenck) C. 



Trevizan Chiomento et al. ‐ Mycorrhizal biotechnology in the goldenberry cultivation

267

Walker & A. Schüßler], arranged in a randomized 
block design with five replications. Each plot consist­
ed of three goldenberry plants. 
     The AMF community used came from the crop­
trap of agricultural soil collected at a reference site 
for strawberry cultivation in the municipality of São 
José do Hortêncio (29° 29’ 33” S, 51° 12’ 24” W), Rio 
Grande do Sul State, Brazil (Chiomento et al., 2019 
a), composed of ten fungal species according to the 
c l a s s i fi c a ti o n  o f  G l o m e r o m y c o t a  p r o p o s e d  b y 
Redecker et al. (2013): Acaulospora foveata Trappe & 
Janos, Claroideoglomus aff. luteum, Claroideoglomus 
claroideum (N.C. Schenck & G.S. Sm.) C. Walker & A. 
Schüßler, Claroideoglomus etunicatum (W.N. Becker 
& Gerd.) C. Walker & A. Schüßler, Funneliformis aff. 
geosporum, Funneliformis aff. mosseae, Funneli‐
formis mosseae (T.H. Nicolson & Gerd.) C. Walker & 
A. Schüßler, Glomus aff. versiforme, Glomus sp. (cae­
saris like) and Glomus sp2. The isolate G. intraradices 
came from the commercial product MYKE® PRO and 
t h e  i s o l a t e  R .  c l a r u s  w a s  o b t a i n e d  f r o m  t h e 

International Collection of Glomeromycota Culture 
(CICG). 
 
Cultivation procedures 
     We applied the treatments (AMF) in two stages: 
1) in the acclimatization of the seedlings; 2) in trans­
planting to the place of cultivation. Thus, of the total 
amount of mycorrhizal inoculant used (10 g), we 
applied 5 g in the acclimatization of the seedlings and 
5 g at the time of transplanting. 
     In September (spring) of 2018, thirty days after 
the germination of goldenberry seeds, the plants 
obtained (Fig. 1D) were acclimatized in 72­cell poly­
styrene trays, filled with the sterilized Horta 2® sub­
strate (120°C for 20 minutes) and with treatments 
related to mycorrhization (1/2 of the total amount), 
with the purpose of seedling production. Horta 2® is 
composed of pine bark, vermiculite, acidity correc­
tors and fertilizers (nitrogen, phosphorus and potas­
sium) in amounts not supplied by the manufacturer. 
A 500 g sample of the substrate was analyzed to 
obtain its physical (Brazil, 2007) and chemical (MAPA, 
2014) attributes (Table 1). 
     The trays were kept on metal benches, 1.2 m from 
the ground surface, in a greenhouse (90 m2), installed 
in the northeast­southeast direction, with a semicir­
cular roof. The galvanized steel structure is covered 
with a low­density polyethylene film with anti­ultravi­
olet additive (150 micron thickness) and the sides are 
covered with an anti­aphid screen. The irrigation 
used during acclimatization was with sprinklers (1.8 
L.min­1 per unit), in the mechanized system. The irri­
gation regime consisted of activating the sprinklers 
seven times a day, with total wetness of 14 minutes. 
The water depth supplied to the seedlings was 7.8 
mm.day­1. 
     In December (summer) 2018, after three months 
of acclimatization, the seedlings were transplanted 

Fig. 1 ­ Obtaining goldenberry plants. (A) Separation of seeds 
from fruits. (B) Selection of seeds. (C) Germination in ger­
box. (D) Plants produced.

Substrate
Physical properties

D (Kg m­3) TP (m3.m­3) AE (m3.m­3) RAW (m3.m­3) BW (m3.m­3) RW (m3.m­3)
Horta 2 ® 241 0.837 0.303 0.149 0.020 0.365

Chemical properties

N % (m.m­1) P2O5 (m.m
­1) K2O % (m.m

­1) OC % (m.m­1) pH EC % (mS.cm­1) CEC (mmolc.kg
­1)

0.36 0.39 0.00 12.60 6.1 0.45 278.60

Table 1 ­ Physical and chemical properties of the Horta 2® substrate

(z) D= density; TP= total porosity; AE= aeration space; RAW= easily available water; BW= buffer water; RW= remaining water. 
(w) N= nitrogen; P2O5 = phosphorus pentoxide; K2O= potassium oxide; OC= organic carbon; pH= hydrogen potential; EC= electric conducti­

vity; CEC= cation exchange capacity. 



Adv. Hort. Sci., 2022 36(4): 265­273

268

into pots (3.6 L), filled with sterilized Horta 2® (120°C 
for 20 minutes) and complemented with the other 
part of the treatments related to mycorrhization (1/2 
of the total amount). The pots were kept in beds cov­
ered with mulching, in a greenhouse (430 m2), with a 
semicircular roof, installed in the northeast­south­
east direction. The galvanized steel structure was 
covered with a low­density polyethylene film (150 
microns thick) and with an anti­ultraviolet additive. 
     Localized irrigation was carried out using drip rods 
(2.4 L.h­1 per unit), in the mechanized system. The 
irrigation regime consisted of activating the dripping 
rods six times a day, with total wetting for six min­
utes. The nutrient solutions supplied to the plants, 
fortnightly, were made according to Furlani and 
Fernandes Júnior (2004), but with a 50% reduction in 
phosphorus supply. Through a mini meteorological 
station, we verified that the average general temper­
ature recorded during the experiment was 25.66°C. 
     The plants were conducted with three stems and 
were tutored with the aid of wires. No biocides were 
used during the crop cycle. The evaluations started 
after the fruiting of the plants, in February (summer) 
of 2019. We evaluated the root system morphology 
and the productive yield (number and weight) and 
quality of fruits. 

Root system morphology 
     At the end of the experiment, in July 2019, the 
plants roots were washed in water to eliminate sub­
strate fragments. The roots were digitized by a scan­
ner and the images obtained were analyzed by the 
WinRHIZO® software. The attributes evaluated were 
total length (TL, cm), surface area (SA, cm2) and vol­
ume (V, cm3). The roots were grouped by the soft­
ware into different diameter classes in relation to 
their total length (Böhm, 1979): very thin (VT, Ø<0.5 
mm), thin (TH, Ø from 0.5 to 2 mm) and thick (TK, 
Ø>2 mm). 
     To verify the infective capacity of AMF, root por­
tions of mycorrhizal plants were prepared according 
to Phillips and Hayman (1970) and their percentage 
of mycorrhizal colonization (MC) was determined 
according to Trouvelot et al. (1986), by the equation: 
 
MC(%) = (total number of fragments with mycorrhizal roots) x 100 
                                        (total number of fragments)                 (1) 
 
Fruit production 
     From fruiting, in February 2019, the total number 
of fruits (TNF, number per plant) and the total pro­

duction of berries (TP, grams per plant) were evaluat­
ed. In addition, the average fresh fruit mass (AFFM, 
grams) was evaluated. The fruits were harvested 
when they were in the stages of maturation between 
4 and 6 (ICONTEC, 1998). The fruits were weighed on 
an electronic digital scale. 
 
Chemical fruit quality 
     The analysis of fruit quality was performed at the 
end of the experiment, in July 2019. The chemical 
characteristics of the fruits were evaluated regarding 
the content of total soluble solids (TSS, %) and total 
titratable acidity (TTA, % of citric acid), from 20 fruits 
of each treatment for each repetition. The TSS con­
tent was determined in an analog refractometer, and 
the TTA was performed according to the norms of 
the Adolfo Lutz Institute (Zenebon et al., 2008). To 
evaluate the flavor of the fruits, the TSS/TTA ratio 
was determined. 
 
Data analysis 
     The data obtained were submitted to analysis of 
variance (Anova) and the averages of the treatments 
were compared by the Tukey test, at 5% error proba­
bility, with the aid of the Costat® program (Cohort 
Software, 2003). 
 
 
3. Results 
 
Root system morphology 
     We verified a significant effect of mycorrhizal 
inoculants only for the attributes MC, V and TH. G. 
intraradices had a greater ability to infect plant roots 
than the mycorrhizal community and R. clarus (Fig. 
2A). The fungal structures identified inside the roots 
of goldenberry plants were hyphae, vesicles and 
arbuscules. In addition, plants inoculated with R. 
clarus had 41% and 42% greater root volume than 
non­mycorrhizal plants and those inoculated with the 
AMF community, respectively (Fig. 2B). Also, plants 
inoculated with the mycorrhizal community had a 
greater amount of fine roots (+47%) compared to the 
control (Fig. 2C). 
 
Fruit production 
     We observed a positive effect of treatments only 
for the AFFM attribute. Non­mycorrhizal plants pro­
duced fruits with higher average fresh mass (+29%) 
compared to plants inoculated with the mycorrhizal 



Trevizan Chiomento et al. ‐ Mycorrhizal biotechnology in the goldenberry cultivation

269

community, but did not differ from plants mycor­
rhizal with G. intraradices and R. clarus (Fig. 3). 

Chemical fruit quality 
     M y c o r r h i z a l  i n o c u l a n t s  i n fl u e n c e d  T S S  a n d 
TSS/TTA attributes (Fig. 4). Sweeter (Fig. 4A) and 
tastier (Fig. 4B) fruits were produced by plants inocu­
lated with the mycorrhizal community and with the 
isolate R. clarus. 
 
 
4. Discussion and Conclusions 
 
     Here, we show that goldenberry plants in the 
absence and presence of AMF inoculation differed in 
horticultural performance. In the first productive 

cycle of the plants, we did not observe the effect of 
mycorrhization on the production of berries. We 
believe that in goldenberry the fungal species used 
require a period of more than one year to benefit the 
fruit yield, as occurs in strawberry (Fragaria X 
ananassa Duch.) cultivation (Robinson­Boyer et al., 
2016). However, goldenberry plants inoculated with 
A M F  s h o w e d  a  m o r e  p r o f u s e  r o o t  s y s t e m . 

Fig. 2 ­ Root system morphology of goldenberry plants in the 
p r e s e n c e  a n d  a b s e n c e  o f  A M F  i n o c u l a t i o n .  ( A ) 
Mycorrhizal colonization (%). (B) Root volume (cm³). (C) 
Amount of fine roots (cm). Data presented as mean ± 
standard deviation. Means followed by the same letter in 
the column did not differ significantly by the Tukey test 
(p≤0.05).

Fig. 3 ­ Average fresh fruit mass (grams) of goldenberry plants in 
the presence and absence of AMF inoculation. Data pre­
sented as mean ± standard deviation. Means followed by 
the same letter in the column did not differ significantly 
by the Tukey test (p≤0.05).

Fig. 4 ­ Chemical quality of goldenberry fruits in the presence 
and absence of AMF inoculation. (A) Total soluble solids 
(%). (B) Fruit flavor. Data presented as mean ± standard 
deviation. Means followed by the same letter in the 
column did not differ significantly by the Tukey test 
(p≤0.05).



(Abbott and Robson, 1981), which limits the plant’s 
response to mycorrhization (Lambais and Cardoso, 
1990). Although mycorrhizal colonization is impor­
tant, the percentage of root infectivity is not always 
c o r r e l a t e d  w i t h  t h e  e ffi c i e n c y  o f  s y m b i o s i s 
(Konvalinková and Jansa, 2016). 
     This lack of relationship between infectivity and 
AMF efficiency in improving crop growth may be 
related to the time required for the establishment of 
root colonization (Abbott and Robson, 1981). Under 
long­term, arbuscular mycorrhiza promotes more 
benefits to the plant host (Ortas, 2012), mainly by 
increasing the acquisition of water and minerals to 
the plant. The major function of AMF is suggested to 
be nutrient acquisition (Zhang et al., 2019) and thus, 
under high nutrient conditions, AMF can shift from a 
net benefit to a cost for the host (Johnson et al., 
2015). In our study, the nutrients supplied during cul­
tivation were not limited to goldenberry; therefore, 
the inoculation effect may not have been potentiated 
in terms of fruit yield. 
     However, we proved the benefit of mycorrhiza­
tion on the chemical quality of the fruits through the 
increase in the sugar content (Fig. 4A) and the better 
berry flavor (Fig. 4B), as already reported for zucchini 
(Cucurbita pepo L.) (Rouphael et al., 2015), strawber­
ry (Costa et al., 2020) and tomato (Solanum lycoper‐
sicum L.) (Sellitto et al., 2019). Plants grown with 
AMF showed a more developed root system (Fig. 2), 
which allows extrarradicial hyphae to extend beyond 
the rhizosphere, making water acquisition more effi­
cient (Xu et al., 2017). Due to greater water availabili­
ty, these plants have greater stomatal opening, 
which increases the rate of transpiration and, thus, 
there is a greater supply of carbon dioxide for photo­
synthesis (Vicente­Sánchez et al., 2014). As a result, 
there is a greater production of sugars, which are the 
primary source of photosynthesis, and this explains 
the increase in the fruit sugar content and, conse­
quently, the best flavor of the berries (Fig. 4). 
     The lack of effect of G. intraradices on berries 
quality can be attributed to the low effectiveness of 
this fungal species on goldenberry. Various factors 
modulate the arbuscular mycorrhiza effect on the 
performance of their associated plants and this 
includes the traits of the host and the fungi them­
selves (Chiomento et al., 2019 a). Cultivated plants 
vary in their responsiveness to AMF due to their mor­
phology (Chiomento et al., 2019 b) and AMF differ in 
the benefits provided to the plant (Werner and Kiers, 
2015). 

270

Adv. Hort. Sci., 2022 36(4): 265­273

Furthermore, the use of the mycorrhizal community 
and R. clarus allowed to harvest fruits with better 
chemical quality. This indicates that the fungal 
species present in these two inoculants have an affin­
ity for this horticultural crop. The use of AMF com­
patible with the host commonly provides more satis­
f a c t o r y  r e s u l t s  ( C h i o m e n t o  e t  a l . ,  2 0 2 2 ) .  G . 
intraradices was more effective in colonizing the 
roots of the plant host. However, the mycorrhizal 
community tested in this study stood out in relation 
to the fungal isolates for generally improving the hor­
ticultural performance of goldenberry. 
     We verified that the root system of the mycor­
rhizal plants was more profuse, with greater volume 
(Fig. 2B) and with a greater amount of fine roots (Fig. 
2C). This benefit to the roots has already been 
reported by Reyes et al. (2019), who demonstrated 
that mycorrhization in goldenberry under water 
stress increased the accumulation of root dry matter. 
Due to the plasticity of the roots, their characteristics 
can be modulated by several factors, including AMF 
(Hodge et al., 2009). During the establishment of the 
association between host and fungus, many molecu­
lar signals are initiated, including an AMF (lipo­chito­
oligosaccharides) diffusible factor called “Myc fac­
tor”, which stimulates the formation of finer roots 
(Oláh et al., 2005), altering the morphology of the 
root system of plants. In addition, root modifications 
under mycorrhization may be related to the alloca­
tion of sugars to roots (Wu et al., 2011) and hormon­
al regulation (Zou et al., 2017), independent of sym­
biotic signaling (Gutjahr, 2014). The more fine roots 
there are in mycorrhizal plants, the better their 
acquisition of water and nutrients (Chiomento et al., 
2021), as these roots are the ones that most acquire 
and use the available resources in the plant growth 
medium (Costa et al., 2019). 
     Differently from what was expected, the produc­
tive performance of goldenberry was higher when 
the plants were not mycorrhized (Fig. 3). This sug­
gests that AMF initially demand carbon from the host 
for their maintenance and only later repay this bene­
fit to the plant symbiont. These results, however, 
contradict the literature. For example, Miranda et al. 
(2011) reported benefits of AMF in the production of 
goldenberry, under limiting conditions to plants, 
which were subjected to abiotic stresses, which did 
not happen in our study. There is not always a high 
relationship between fungal infectivity and efficiency 
in promoting crop growth due to the time required to 
establish bidirectional flow between symbionts 



Trevizan Chiomento et al. ‐ Mycorrhizal biotechnology in the goldenberry cultivation

271

     The two genera that made up the monospecific 
inoculants, Glomus and Rhizophagus, have already 
been reported in studies of AMF diversity in golden­
berry (Ramírez­Gómez et al., 2019) and have also 
been used in applicability studies (Rhizophagus) in 
t h i s  h o r ti c u l t u r a l  c r o p  ( M i r a n d a  e t  a l . ,  2 0 1 1 ) . 
However, in our study, the mycorrhizal community, 
representing a multispecific inoculant, stood out in 
relation to the fungal isolates for generally improving 
the horticultural performance of goldenberry, mainly 
by increasing the amount of fine roots produced by 
the plants (Fig. 2C) and for benefiting fruit quality 
(Fig. 4). Inoculation with AMF populations, such as 
the fungal community used in this study, generally 
provides more satisfactory results due to greater 
compatibilities at the fungus­host interface and by 
increasing mutualistic effects with two or more sym­
bionts instead of just one (Chiomento et al., 2019 b). 
This mycorrhizal community was obtained through 
the trap culture technique, that is, produced on‐
farm. 
     A mycorrhizal inoculant rich in propagules and 
produced on‐farm may be a suitable solution for 
large­scale inoculation of crops (agroecosystems), 
seedlings (nurseries) and in potting media for veg­
etable growers (soilless cultivation) (Douds Junior et 
al., 2006). The use of non­sterilized growth medium 
(soil and/or substrate) as a component of the on­
farm inoculant represents a source of propagules for 
other AMF present in this growth medium, which 
results in an inoculant with greater taxonomic diver­
sity (Schlemper and Stürmer, 2014). This is strongly 
desired due to the functional diversity exhibited by 
mycorrhizal species for the promotion of plant 
growth, for example (Chiomento et al., 2020 b). 
Furthermore, a diversified inoculant potentiates a 
combination between fungal isolates and an eventual 
host (Douds Junior et al., 2006). 
     Therefore, the results of our research confirmed 
the potential of applying mycorrhizal biotechnology 
to goldenberry, as the work demonstrated that AMF 
can be a valuable tool for the cultivation of this veg­
etable. The complex roles of AMF in agroecosystems 
are just beginning to be understood (Chiomento et 
al., 2022). In this way, a greater understanding of the 
application and benefits of AMF can enable their use 
in the sustainable production of vegetables (Trentin 
et al., 2022). 
     We conclude that in the first production cycle, 
there is no effect of mycorrhization on the total num­
ber of fruits and total production of berries. We 

believe that for the goldenberry cultivation the fun­
gal species used require a period of more than one 
year to benefit the fruit yield. On the other hand, 
goldenberry plants submitted to mycorrhizal biotech­
nology have a more profuse root system and produce 
fruits with better chemical quality. G. intraradices is 
most effective in colonizing the plant host roots. 
However, the mycorrhizal community stands out in 
relation to the fungal isolates for generally improving 
the horticultural performance of goldenberry. Thus, 
the application of this biotechnological tool in the 
goldenberry culture can be an alternative to spread 
and promote its sustainable cultivation. 
 
 
Acknowledgements 
 
     AMF used in this work are regulated by Sistema 
Nacional de Gestão do Patrimônio Genético e do 
Conhecimento Tradicional Associado (SisGen) of the 
Ministry of the Environment, Brazil, according to the 
registration number A198F50. 
 
 
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