© The Authors, 2023, Published by the Universidad del Zulia*Corresponding author: evelyncpp@gmail.com
Keywords:
Phenology
Environment
Secondary metabolites
Phenology and environment in the presence of secondary metabolites in Psidium guajava L.
Fenología y ambiente en la presencia de metabolitos secundarios en Psidium guajava L.
Fenologia e ambiente na ocorrência de metabólitos secundários em Psidium guajava L.
Evelyn del Carmen Pérez Pérez
1
*
Gretty Rosario Ettiene Rojas
2
Maribel del Carmen Ramírez Villalobos
3
Ángel Gómez Degraves
4
Rev. Fac. Agron. (LUZ). 2023, 40(4): e2340Spl04
ISSN 2477-9407
DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v40.supl.04
Crop production
Associate editor: Dr. Jorge Vilchez-Perozo
University of Zulia, Faculty of Agronomy
Bolivarian Republic of Venezuela
Abstract
Guava (Psidium guajava L.) is one of the most important fruit trees in
Venezuela due to the acceptance of its fresh and processed fruit with relevant
sensory and nutritional characteristics. The establishment of the crop in the
producing areas has been the result of the initiative of leading farmers in the
country. Due to the potential of guava, the present review aimed to describe
the phenology, environment, and presence of secondary metabolites in P.
guajava. The search for information on P. guajava included several key
words such as phenology, owering, fruiting, and secondary metabolism.
Sixty-six references were selected from 130 results, including research
articles, reviews, and books published between 1991 and 2023. Secondary
metabolite biosynthesis is a dynamic process that depends on numerous
factors associated with the plant and the environment. The content of phenols
and avonoids in guava can help characterize its production, agroindustrial,
and pharmaceutical importance, be a tool for cultivar selection, and anticipate
the content of other secondary metabolites to identify plants that dier in
their production.
1
Departamento
de
Agronomía.
Facultad
de
Agronomía.
Universidad del Zulia, Maracaibo, Zu 4005. Venezuela.
2
Departamento
de
Química.
Facultad
de
Agronomía.
Universidad del Zulia, Maracaibo, Zu 4005. Venezuela.
3
Departamento
de
Botánica.
Facultad
de
Agronomía.
Universidad del Zulia, Maracaibo, Zu 4005. Venezuela.
4
Departamento
de
Estadística.
Facultad
de
Agronomía.
Universidad del Zulia, Maracaibo, Zu 4005. Venezuela.
Received: 09-10-2023
Accepted: 22-11-2023
Published: 06-12-2023
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2023, 40 (Supplement): e2340Spl04. October-December. ISSN 2477-9407.2-6 |
Resumen
El guayabo (Psidium guajava L.) se ubica entre los frutales
más relevantes en Venezuela por la aceptación de su fruta fresca y
procesada con características sensoriales y nutricionales relevantes.
El establecimiento del cultivo en las zonas productoras se ha
realizado por la iniciativa de agricultores líderes del país. Debido
al potencial del guayabo la presente revisión tuvo como objetivo
describir la fenología, el ambiente y la presencia de metabolitos
secundarios en P. guajava. La búsqueda de información sobre P.
guajava incluyó varias palabras clave como fenología, oración,
fructicación y metabolismo secundario. Se seleccionaron 66
referencias de 130 resultados, incluyendo artículos de investigación,
revisiones y libros publicados entre 1991 y 2023. La biosíntesis de
metabolitos secundarios es un proceso dinámico que depende de
numerosos factores asociados a la planta y al ambiente. El contenido
de fenoles y avonoides en guayabo puede ayudar a caracterizar
la producción e importancia agroindustrial y farmacéutica, ser una
herramienta para la selección de cultivares y anticipar el contenido
de otros metabolitos secundarios para identicar plantas que se
diferencien en la producción de ellos.
Palabras clave: fenología, ambiente, metabolitos secundarios.
Resumo
A goiaba (Psidium guajava L.) é uma das árvores frutíferas mais
importantes na Venezuela devido à aceitação de seus frutos frescos e
processados com características sensoriais e nutricionais relevantes.
O estabelecimento da cultura nas áreas produtoras tem sido efectuado
por iniciativa dos principais agricultores do país. Devido ao potencial
da goiaba, a presente revisão teve como objetivo descrever a fenologia,
o ambiente e a presença de metabólitos secundários em P. guajava. A
busca de informações sobre P. guajava incluiu várias palavras-chave,
como fenologia, oração, fruticação e metabolismo secundário.
Foram seleccionadas sessenta e seis referências de 130 resultados,
incluindo artigos de investigação, revisões e livros publicados entre
1991 e 2023. A biossíntese de metabolitos secundários é um processo
dinâmico que depende de numerosos factores associados à planta e ao
ambiente. O conteúdo de fenóis e avonóides na goiaba pode ajudar a
caraterizar a produção e a importância agroindustrial e farmacêutica,
ser uma ferramenta para a seleção de cultivares e antecipar o conteúdo
de outros metabolitos secundários para identicar plantas que diferem
na sua produção.
Palavras-chave: fenologia, ambiente, metabólitos secundários.
Introduction
Guava (Psidium guajava L.; family: Myrtaceae) cultivation in
Venezuela faces challenges due to oversupply and high costs derived
from the excess of chemical inputs for pest, disease and weed
management. This causes a decrease in market prices and protability,
which hinders the international positioning of the crop. Guava fruit
has excellent values in minerals, vitamins (Vijaya et al., 2020) and
secondary metabolites such as phenols and avonoids (Pérez-Pérez
et al., 2014; 2019; 2020) generating biotechnological interest for
its antioxidant, anti-inammatory, antibacterial, antihyperlipidemic,
cardioprotective, antimutagenic (Naseer et al., 2018), antifungal
(Liu et al., 2018), antidiabetic, hepatoprotective (Li et al., 2021) and
larvicidal (Mendes et al., 2017) properties.
The leaves contain phenols (Pérez-Pérez et al., 2014) and
avonoids (Pérez-Pérez et al., 2014; Rivero-Maldonado et al., 2013),
and its essential oil is rich in phenolic compounds from the tannin and
avonoid groups (Jassal and Kaushal, 2019). Secondary metabolite
biosynthesis is a dynamic process inuenced by factors such as plant
type and ontogenetic stage (Verma and Shukla, 2015). The variety of
compounds in plants supports phenotypic plasticity (Kaplan et al.,
2008).
The aforementioned allows describing the relevance of P. guajava
plants in the present review, in correspondence with phenology, the
relationships between the environment and phenology, phenological
phases and the presence of secondary metabolites and between the
environment and the presence of secondary metabolites, this based
on Verma and Shukla (2015), who consider that the biosynthesis of
secondary metabolites is a dynamic process that depends mainly
on numerous factors associated with the plant itself and with the
environment. Within the rst group of factors, the most important
are the type of plant and the stage of its ontogenetic development.
The objective of this review was to describe the phenology, the
environment and the presence of secondary metabolites in P. guajava.
Methods
The search words or phrase were “Psidium guajava L.”,
“phenology”, “owering”, “fruiting”, “vegetative sprouting”,
“secondary metabolites”, “phenolic compounds”, “avonoids”,
“antioxidant capacity”, “biotic stress” and abiotic stress”. Scientic
literature was searched using the web search engine Scholar Google,
Scopus, Web of Science, Taylor and Francis, Springer Link, PubMed
and Science Direct. For the review 66 references were selected from
the 130 resulting from the search; those included research articles,
review articles and books published between 1991 and 2023.
Discussion
Phenology of P. guajava
Phenology included owering, fruiting, budding and leaf fall on a
temporal scale to identify the vegetative and reproductive processes
of the species (Biondi et al., 2007). These vary among genotype,
edaphoclimatic conditions and crop management (Fisher and Orduz-
Rodriguez, 2012). The adult guava plant has been described as a semi-
deciduous tree, because after harvesting it undergoes a phenomenon
of exhaustion or lethargy with the presence of yellowing and the fall
of most of the leaves, which continues during the dry period, resuming
the growth of new branches and bud regrowth with the onset of the
rainy season (Gómez, 1995).
This phenological development was evidenced in the municipality
of Mara, Zulia state, Venezuela (Marín et al., 2000); the study showed
that it was mainly due to the interaction of age and/or genetic potential
with the environment. Pérez-Pérez (2023) armed the occurrence
of the phenological phases of fruiting, vegetative sprouting and
owering, the rst being the most frequent, followed by vegetative
sprouting, a behavior that corresponded with that reported by Esparza
et al. (1993) and Marín et al. (2004). In turn, Marín et al. (2000)
identied fruit set in all plants throughout the phenological stages
of vegetative and reproductive sprouting, during the evaluation of P.
guajava Criolla roja, Cubana and Montalbán cultivars grafted on Cas
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3-6 |
In the case of tropical conditions, phenological changes are dened
by periods of drought and rain, manifesting physiological changes
with increased growth, swelling and bud detachment, increased
stem diameter, oral initiation and dierentiation, initiation of fruit
set and fruit ripening (Quirós et al., 2009; Salazar et al., 2006), with
the dierence that guava can produce fruit throughout the year with
maximum and minimum production peaks, depending on climatic
conditions (Marin et al., 2000; 2004; Pérez-Pérez et al., 2023).
Marín et al. (2000), in their study on the behavior of guava types
grafted on P. friedrichsthalianum Berg-Niedenzu, reported vegetative
and reproductive sprouting with the presence of physiologically
mature fruit on all plants. This is due to the behavior of this species,
which maintains the three phenological stages active on the plant
throughout the year (Marín et al., 2004), with a variation in the
intensity of occurrence, responding to the climate in the area.
Pérez-Pérez et al. (2023) studied the environmental impact on
guava ower, fruit and vegetative bud production in Mara, Zulia,
Venezuela. These researchers emphasized the highest production of
vegetative buds in October-November, the subsequent decrease and
the maximum in may. Precipitation aected the vegetative budburst
phase (da Fontoura et al., 2021) between October and February,
while absence in March aected it, similar to Marín et al. (2000).
The researchers did not identify any signicant relationship with
temperature, humidity, evaporation and insolation (Mendoza et al., 2017).
Flower production alternates with fruit production. The higher or
lower occurrence of fruit corresponds to the lower or higher ower
production that occurred 90 days earlier (Pérez-Pérez et al., 2023;
Singh, 2011), as a response to the higher amount of water available to
the plants due to the occurrence of rainfall (Marín et al., 2000).
Relationship between the phenological phases of P. guajava and
the presence of secondary metabolites.
In the course of evolution, plants have developed the ability to
produce an enormous amount of phenolic secondary metabolites,
which are not necessary in the primary processes of growth and
development, but are of vital importance for their interaction with the
environment, for their defense mechanisms (Cheynier et al., 2013)
and perpetuate their species (Lustre, 2022; Mamani and Filippone,
2018; Rioja Soto, 2020).
A large number of secondary molecules are biosynthesized from
primary metabolites and accumulate in plant cells (Böttger et al.,
2018; Narayani et al., 2017; Rejeb et al., 2014), whose production
could be induced under in vitro conditions when cell cultures by
using biotic, abiotic elicitors and signaling molecules and are usually
increased by specic stress eect (Piñol et al., 2013).
The accumulation and concentration of polyphenolic compounds
varies greatly in dierent plant parts and organs, closely related to
their function in the life cycle and growth phase (Verma and Shukla,
2015). For example, during the fruiting phase of guava, leaves have
the highest amount of phenols, compared to vegetative shoots and
owers that contain less (Pérez-Pérez et al., 2023).
Phenols can modulate essential physiological processes, such as
transcriptional regulation and signal transduction. Some interesting
eects of phenols in plants are also those associated with growth
hormone (auxin). An additional role has been observed for avonoids
in the functional development of pollen. Finally, anthocyanins
represent a class of avonoids that provide the orange, red and
blue/purple colors to many plant tissues. According to co-evolution
theory, red is a signal of tree status to insects migrating to (or moving
between) trees in autumn (Cheynier, 2013).
(P. friedrichsthalianum). They also pointed out that the dierences
in phenological behavior were aected by the interaction between
canopy/pattern and environment, reporting that owering time varied
among the materials from 15 to 45 days.
Marín et al. (2004) evaluated organic amendments in the recovery
of guava trees growing in a eld infested with nematodes of the genus
Meloidogyne, in the state of Zulia, Venezuela. The study showed
that during the evaluation period, the plants that received organic
amendments registered continuous vegetative and reproductive
structures, with the highest fruiting occurring during the month of
July. According to Esparza et al. (1993), this behavior is due to the
physiological response of the plant to the degree of soil moisture
that promotes the onset of the owering phase, which is directly
associated with the fruiting potential, nding a greater supply of fruit
in the months from June to August (50.6 %) and from November to
January (31.1 %) remaining production.
Tong et al. (1991) documented guava production in Mara,
Venezuela, in two seasons, each inuenced by the bimodal rainfall of
the area, with signicant ower emergence. However, in P. guajava
the phenological phases overlap (Pérez-Pérez et al., 2023), due to the
continuity of guava production during the year (Marín et al., 2000).
The growth of vegetative buds during the owering and vegetative
sprouting phase (Pérez-Pérez et al., 2023) has been associated with the
preparation of vegetative organs, mainly leaves, which will guarantee
the production of photoassimilates, such as carbohydrates, that will
support the development of reproductive structures, including fruits.
In general, the guava plant takes about 100 to 150 days from owering
to harvest (Singh, 2011).
In this regard, Pawar and Rana (2019) have pointed out that the
balance between vegetative and reproductive growth is an important
aspect to improve the yield and quality of fruit trees, based on the
source-sink relationship, the authors stated that profuse owering and
fruiting create a high demand for the limited source of carbohydrates,
which aects fruit set and fruit development, hence the importance
of knowing the regulation of carbon distribution in fruit plants in the
short and long term, given the perennial nature of the trees.
Relationship between environment and phenology of P.
guajava
The growth cycle depends on the genotype of the plant (Salazar
et al., 2006) and the environment. Hence, it is essential to evaluate
the climatic circumstances that favor plant development (Ferreira
et al., 2019). When grown in dierent environments, identical
genotypes can present multiple developmental stages. Hence, it
is essential to understand the phenological cycles of crops in order
to dene agronomic management strategies in accordance with the
phenological stages of the plant (Salazar et al., 2006).
Another aspect to consider, during owering and fruiting of guava,
is that environmental conditions and nutrition play an important role
in their determination. For ecient owering and fruiting, sunlight
is necessary; however, periods of water stress are critical to improve
owering. Often under favorable conditions, many of the owers
emerging from the leaf axils of young shoots (solitary or in groups
of two or three), generate fruit on trees (Shivpoojan et al., 2018). In
this regard, Mendoza et al. (2017) noted that the reproductive phase
of plants was promoted by precipitation by 73.4 %, compared to
temperature (19.3 %) followed by solar radiation or photoperiod (3.2
%), results obtained in their study on agrometeorological inuence on
the reproductive phase.
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Rev. Fac. Agron. (LUZ). 2023, 40 (Supplement): e2340Spl04. October-December. ISSN 2477-9407.4-6 |
Phenols are synthesized in leaves (Vranová et al., 2012),
participate in their development (Pawar and Rana, 2019) and from
there are transported to other tissues and organs. Leaves have higher
concentrations of these compounds than the rest of the plant (Vranová
et al., 2012). However, they are not uniformly found throughout the
plant and are often restricted to particular organs and to certain cells
and tissues within that organ (Vranová et al., 2012).
The leaf is the main organ where photosynthesis takes place
and provides carbohydrates that are diverted to the reproductive
structures (Fotirić et al., 2020; Pawar and Rana, 2019) and to the
root apices and stems of the plant (Pawar and Rana, 2019). The fate
of carbohydrates formed during photosynthesis, part of which will
be used in the structural makeup of the plant (primary metabolism)
(Azad et al., 2020) and others in secondary metabolism, will have a
major inuence on the quantity and quality of phenolic compounds
produced (Saltveit, 2017).
Flavonoids play important roles in plant biology, responding to
light and controlling the levels of growth-regulating auxins, plant
dierentiation and in virtually every interaction a plant establishes
with its environment (Vicente and Boscaiu, 2018). Among these,
anthocyanins (Peñarrieta et al., 2014), act in the development of petal
color to attract pollinators (Zhang et al., 2014).
Flavonoids (anthocyanins, avones, avonols and isoavones)
are the most abundant phenolic compounds, have an important
role in plant growth, as well as in the defense mechanism against
microorganisms and insects (Shen et al., 2022) and are abundant in
woody species (Baskar et al., 2018). In the leaf, they are found in
high concentration as catechin and epicatechin (Pérez-Pérez et al.,
2020). The highest avonoid content is found in the owering phase,
followed by the vegetative sprouting and fruiting phase (Pérez-Pérez
et al., 2023), due to the competition for carbon skeletons between
avonoid synthesis and the synthesis of compounds involved in fruit
cell division (Li et al., 2021).
Guava leaf extracts contain phenolic compounds such as gallic
acid, catechin, chlorogenic acid, caeic acid, epicatechin, rutin,
quercetin, kaempferol and luteolin, crucial for antioxidant activity,
not only because of their ability to donate hydrogen, but also because
of their stable intermediate radicals, which prevent oxidation (Morais-
Braga et al., 2017).
Determinations carried out in dierent geographical locations
have reported compounds such as avonoids, glycosides and
aglycones (Wang et al., 2017). As asserted by Camarena-Tello et al.
(2018), because phenolic compounds are considered the main group
of secondary metabolite present in plants (Tolić et al., 2017).
The guava plant directs its secondary metabolism towards the
production of metabolites at the level of various plant organs such as
the leaf (Azad et al., 2020), which are being demanded by the organs
involved in the phenological phase in which the plant is found (Verma
and Shukla, 2015). Thus, higher ower production was associated with
higher production of avonoids determined in the leaf (Pérez-Pérez
et al., 2023). For Coutinho (2013), secondary metabolites represent
a chemical interface between plants and the environment, therefore,
their synthesis is often aected by environmental conditions.
Eect of the environment on the presence of secondary
metabolites
Plants produce secondary metabolites during growth, which serve
a variety of cellular essential for physiological processes and stress
response signaling (Isah, 2019). The type and concentration of these
molecules depend on the species, genotype, developmental stage,
plant tissue or organ, and developmental environment (Isah, 2019;
Speed et al., 2015). Genetic characteristics inuence intrapopulation
variability (Speed et al., 2015), with phenotypic character being
quantied (Arbona et al., 2013), inuenced by genetic and
environmental factors (Turner et al., 2016).
Environmental factors
Environmental factors, including biotic and abiotic stresses,
inuence plant secondary metabolite synthesis. Factors such as
nutrient supply, air temperature, light and water can aect their
concentration (Arbona et al., 2013; Isah, 2019; Speed et al., 2015),
to counteract stress (Arbona et al., 2013; Lustre, 2022; Mamani and
Filippone, 2018). Then environmental factors are decisive in the
biosynthesis of secondary metabolites (Yang et al., 2018).
Appiah et al. (2022) suggest that factors such as development,
seasonality, precipitation, temperature and altitude are correlated
and can inuence secondary metabolism. According to Huyskens-
Keil et al. (2020), high temperatures and plant age can increase
the accumulation of phenolic compounds, while light enhances the
biosynthesis of these compounds. Both biotic and abiotic stresses
stimulate carbon uxes from primary to secondary metabolic
pathways, as noted by Lattanzio (2013).
Biotic factors
Plants are physically attacked by many biological agents such
as fungi, viruses, bacteria, nematodes, among others, which causes
stress in plants known as biotic stress. According to Jan et al. (2021),
plants show resistance against these pathogens through secondary
metabolites. Some secondary metabolites have antimicrobial activities
that function as a plant defensive system against these pathogens (Xu
et al., 2022). The requirement of high concentrations of secondary
metabolites, by plants, during defense against pathogens causes their
biosynthesis to be rapidly triggered.
Abiotic factors
During ontogeny, plants interact with the surrounding environment
and come into contact with dierent abiotic components such as
water, light, temperature, soil, and chemicals (minerals/fertilizers)
(Jan et al., 2021; Xu et al., 2022). Plants require these components
for their development and survival in adequate amounts. However,
more or less of these abiotic components cause stress to plants and
ultimately lead to variations in the production or accumulation of
secondary metabolites during dierent developmental stages of the
plant life cycle, depending on their need (Verma and Shukla, 2015).
Secondary metabolites play an important role in plant adaptation
to the environment and in overcoming stress conditions (Lattanzio,
2013) and consequently plant adaptation to a given area (Pant et al.,
2021).
According to Coutinho (2013) and Sampaio et al. (2016)
temporal and spatial variations, total content, as well as proportions
of secondary metabolites in plants occur at dierent levels (seasonal
and daily, intraplant, inter- and intraspecic) and despite the existence
of genetic control, expression may undergo modications resulting
from the interaction of biochemical, physiological, ecological
and evolutionary processes. They represent a chemical interface
between plants and the surrounding environment, so their synthesis is
frequently aected by environmental conditions (Chiveu et al., 2019).
In a recent investigation Pérez-Pérez et al. (2023) obtained higher
concentrations of avonoids in the fruiting phase in the absence of
precipitation, as well as high temperatures and insolation in the study
area. Espinosa-Leal et al. (2021) and Piasecka et al. (2017) indicated
that plant exposure to drought promoted increased production of
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Pérez et al. Rev. Fac. Agron. (LUZ). 2023, 40 (Supplement): e2340Spl04
5-6 |
several classes of secondary metabolites including terpenes, complex
phenols, and alkaloids during in vitro and in vivo growth through the
induction of ionic or osmotic stress.
Conclusion
Guava is a plant rich in secondary metabolites, particularly
phenolic compounds, such as phenols and avonoids, with biological
anti-inammatory, antimicrobial and antioxidant properties. Scientic
papers on the phenology of P. guajava were found less frequently
than expected. However, it is a characteristic that deserves attention
because it allows the development of agronomic management
techniques to support crop production. Scientic studies have shown
that P. guajava is a highly productive species of secondary metabolites
under stress conditions. The production of secondary metabolites of
the plant could be a useful indicator for the characterization of its
production of agroindustrial and pharmaceutical importance and,
therefore, would constitute a useful tool in the selection process.
Literatura citada
Appiah, K. S., Omari, R. A., Onwona-Agyeman, S., Amoatey, C. A., Ofosu-
Anim, J., Smaoui, A., Arfa, A. B., Suzuki, Y., Oikawa, Y., Okazaki, S.,
Katsura, K., Isoda, H., Kawada, K., & Fujii, Y. (2022). Seasonal changes
in the plant growth-inhibitory eects of rosemary leaves on lettuce
seedlings. Plants, 11(5), 673. Doi: 10.3390/plants11050673
Arbona, V., Manzi, M., de Ollas, C., & Gómez-Cadenas, A. (2013). Metabolomics
as a tool to investigate abiotic stress tolerance in plants. International
Journal of Molecular Sciences, 14(3), 4885-4911. Doi: 10.3390/
ijms14034885
Azad, M. O. K., Kjaer, K. H., Adnan, M., Naznin, M. T., Lim, J. D., Sung, I. J.,
Park, C. H., & Lim, Y. S. (2020). The evaluation of growth performance,
photosynthetic capacity, and primary and secondary metabolite content
of leaf lettuce grown under limited irradiation of blue and red led
light in an urban plant factory. Agriculture, 10(2), 28. Doi: 10.3390/
agriculture10020028
Baskar, V., Venkatesh, R., & Ramalingam, S. (2018). Flavonoids (Antioxidants
Systems) in higher plants and their response to stresses. In: D. Gupta, J.
Palma, F. Corpas (Eds.), Antioxidants and Antioxidant Enzymes in Higher
Plants (pp. 253-268). Springer, Cham. Doi: 10.1007/978-3-319-75088-
0_12
Biondi, D., Leal, L., & Batista, A. C. (2007). Fenologia do orescimento
e fruticação de espécies nativas dos Campos. Acta Scientiarum.
Biological Sciences, 29(3), 269-276. https://www.redalyc.org/articulo.
oa?id=187115762005
Böttger, A., Vothknecht, U., Bolle, C., & Wolf, A. (2018). Plant secondary
metabolites and their general function in plants. In: Lessons on Caeine,
Cannabis & Co: Plant-derived drugs and their interaction with human
receptors. Learning Materials in Biosciences (pp. 3-17). Springer, Cham.
Doi: 10.1007/978-3-319-99546-5_1
Camarena-Tello, J., Martínez-Flores, H., Garnica-Romo, M., Padilla-Ramírez, J.,
Saavedra-Molina, A., Alvarez-Cortes, O., Bartolomé-Camacho, M., &
Rodiles-López, J. (2018). Quantication of phenolic compounds and in
vitro radical scavenging abilities with leaf extracts from two varieties of
Psidium guajava L. Antioxidants, 7(3), 34. Doi: 10.3390/antiox7030034
Cheynier, V., Comte, G., Davies, K. M., Lattanzio, V., & Martens, S. (2013).
Plant phenolics: recent advances on their biosynthesis, genetics, and
ecophysiology. Plant Physiology and Biochemistry, 72, 1-20. Doi:
10.1016/j.plaphy.2013.05.009
Chiveu, J., Naumann, M., Kehlenbeck, K., & Pawelzik, E. (2019). Variation in
fruit chemical and mineral composition of Kenyan guava (Psidium
guajava L.): Inferences from climatic conditions, and fruit morphological
traits. Journal of Applied Botany and Food Quality, 92, 151-159. Doi:
10.5073/JABFQ.2019.092.021
Coutinho, A. (2013). Extração de tanino em folhas, sementes e frutos
verdes de cinamomo (Melia azedarach L.) com diferentes tipos de
solventes (Bachelor’s thesis, Universidade Tecnológica Federal do
Paraná). http://repositorio.utfpr.edu.br/jspui/handle/1/6501
da Fontoura Custódio Monteiro, V., Dias Gonçalves, E., Abreu Moura, P. H.,
Vieira da Silva, L., Bolzan Martins, F., & Norberto, P. M. (2021). Estágios
fenológicos da goiabeira ‘Paluma’ em região de clima subtropical de acordo
com a escala BBCH. Revista Brasileira De Ciências Agrárias, 16(3), 1-8.
Doi: 10.5039/agraria.v16i3a177
Esparza, D., Tong, F., Parra, G., Sosa, L., & Petit, D. (1993). Caracterización de la
producción de guayaba, Psidium guajava L., en una granja del Municipio
Mara del Estado Zulia. Revista de la Facultad de Agronomía de la
Universidad del Zulia, 10(Suplemento 1), 53-54.
Espinosa-Leal, C. A., Mora-Vásquez, S., Puente-Garza, C. A., Alvarez-Sosa, D.
S., & García-Lara, S. (2022). Recent advances on the use of abiotic stress
(water, UV radiation, atmospheric gases, and temperature stress) for the
enhanced production of secondary metabolites on in vitro plant tissue
culture. Plant Growth Regulation, 97(1), 1-20. Doi: 10.1007/s10725-022-
00810-3
Ferreira, M. D. C., Martins, F. B., Florêncio, G. W., & Pasin, L. A. (2019). Cardinal
temperatures and modeling of vegetative development in guava. Revista
Brasileira de Engenharia Agrícola e Ambiental, 23(11), 819-825. Doi:
10.1590/1807-1929/agriambi.v23n11p819-825
Fisher, G., & Orduz-Rodríguez, O. (2012). Ecosiología en frutales. En: G. Fisher,
L.M. Melgarejo, D. Miranda (Eds.). Manual para el cultivo de frutales en
el trópico. (pp. 54-72). Produmedios.
Fotirić, M. F., Tosti, T., Sredojević, M., Milivojević, J., Meland, M., & Natić,
M. (2019). Comparison of sugar prole between leaves and fruits of
blueberry and strawberry cultivars grown in organic and integrated
production system. Plants, 8(7), 205. Doi: 10.3390/plants8070205
Gómez, R. (1995). Manejo agronómico del cultivo del guayabo en Colombia (No.
Doc. 24646) CO-BAC, Bogotá.
Huyskens-Keil, S., Eichholz-Düdar, L., Hassenberg, K., & Herppich, W.B. (2020).
Impact of light quality (White, red, blue light and UV-C irradiation) on
changes in antocyanin content and dynamics of PAL and POD activities
in apical and basal spear sections of White asparagus after harvest.
Postharvest Biology and Thechnology, 161, 111069. Doi: 10.1016/j.
postharvbio.2019.111069
Isah, T. (2019). Stress and defense responses in plant secondary metabolites
production. Biological Research, 52(39), 1-25. Doi: 10.1186/s40659-
019-0246-3
Jan, R., Asaf, S., Numan, M., Lubna, & Kim, K. M. (2021). Plant secondary
metabolite biosynthesis and transcriptional regulation in response to
biotic and abiotic stress conditions. Agronomy, 11(5), 968. Doi: 10.3390/
agronomy11050968
Jassal, K., & Kaushal, S. (2019). Phytochemical and antioxidant screening
of guava (Psidium guajava) leaf essential oil. Agricultural Research
Journal, 56(3), 528-533. Doi: 10.5958/2395-146X.2019.00082.6
Kaplan, I., Halitschke, R., Kessler, A., Sardanelli, S., & Denno, R. F. (2008).
Constitutive and induced defenses to herbivory in above‐and belowground
plant tissues. Ecology, 89(2), 392-406. Doi: 10.1890/07-0471.1
Lattanzio, V. (2013). Phenolic Compounds: Introduction. In: K. Ramawat, J.
Mérillon (Eds.), Natural Products (pp. 1543-1580). Springer. Doi:
10.1007/978-3-642-22144-6_57
Li, X., Li, B., Min, D., Ji, N., Zhang, X., Li, F., & Zheng, Y. (2021).
Transcriptomic analysis reveals key genes associated with the
biosynthesis regulation of phenolics in fresh-cut pitaya fruit (Hylocereus
undatus). Postharvest Biology and Technology, 181, 111684. Doi:
10.1016/j.postharvbio.2021.111684
Li, Y., Xu, J., Li, D., Ma, H., Mu, Y., Zheng, D., Huang, X., & Li, L. (2021).
Chemical characterization and hepatoprotective eects of a standardized
triterpenoid-enriched guava leaf extract. Journal of Agricultural and
Food Chemistry, 69(12), 3626-3637. Doi: 10.1021/acs.jafc.0c07125
Liu, X., Yan, X., Bi, J., Liu, J., Zhou, M., Wu, X., & Chen, Q. (2018). Determination
of phenolic compounds and antioxidant activities from peel, esh, seed of
guava (Psidium guajava L.). Electrophoresis, 39(13), 1654-1662. Doi:
10.1002/elps.201700479
Lustre, H. (2022). Los superpoderes de las plantas: los metabolitos secundarios
en su adaptación y defensa. Revista Digital Universitaria, 23(2). Doi:
10.22201/cuaieed.16076079e.2022.23.2.10
Mamani de Marchese, A., & Filippone, M. P. (2018). Bioinsumos: componentes
claves de una agricultura sostenible. Revista agronómica del noroeste
argentino, 38(1), 9-21. https://ranar.faz.unt.edu.ar/index.php/ranar/
article/view/36/29
Marín, M., Casassa, A., Pérez, E., González, C., Chirinos, D., González, C., &
Sandoval, L. (2004). Enmiendas orgánicas para la recuperación de
árboles de guayabo (Psidium guajava L.) infestados con Meloidogyne
incognita. I. Variación de características fenológicas. Revista de la
Facultad de Agronomía de la Universidad del Zulia, 21(Supl. 1), 129-
136. https://www.produccioncienticaluz.org/index.php/agronomia/
article/view/26529
Marín, M., Casassa, A., Rincón, A., Labarca, J., Hernández, Y., Gómez, E., Viloria,
Z., Bracho, B., & Martínez, J. (2000). Comportamiento de tipos de guayabo
(Psidium guajava L.) injertados sobre Psidium friedrichsthalianum Berg-
Niedenzu. Revista de la Facultad de Agronomía de la Universidad del
Zulia, 17(5), 384-392. https://www.produccioncienticaluz.org/index.
php/agronomia/article/view/26369
Mendes, L. A., Martins, G. F., Valbon, W. R., de Souza, T. D. S., Menini, L.,
Ferreira, A., & da Silva Ferreira, M. F. (2017). Larvicidal eect of essential
oils from Brazilian cultivars of guava on Aedes aegypti L. Industrial
Crops and Products, 108, 684-689. Doi: 10.1016/j.indcrop.2017.07.034
Mendoza, I., Peres, C., & Morellato, L. (2017). Continental-scale patterns and
climatic drivers of fruiting phenology: A quantitative neotropical
review. Global and Planetary Change, 148, 227-241. Doi: 10.1016/j.
gloplacha.2016.12.001
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Pérez et al. Rev. Fac. Agron. (LUZ). 2023, 40 (Supplement): e2340Spl04
6-6 |
Morais-Braga, M., Carneiro, J., Machado, A., Sales, D., dos Santos, A., Boligon,
A., Athayde, M., Menezes, I., Souza, D., Costa, J., & Coutinho, H.
(2017). Phenolic composition and medicinal usage of Psidium guajava
Linn.: Antifungal activity or inhibition of virulence?. Saudi Journal of
Biological Sciences, 24(2), 302-313. Doi: 10.1016/j.sjbs.2015.09.028
Narayani, M., & Srivastava, S. (2017). Elicitation: a stimulation of stress in in
vitro plant cell/tissue cultures for enhancement of secondary metabolite
production. Phytochemistry Reviews, 16(6), 1227–1252. Doi: 10.1007/
s11101-017-9534-0
Naseer, S., Hussain, S., Naeem, N., Pervaiz M., & Rahman, M. (2018). The
phytochemistry and medicinal value of Psidium guajava (guava). Clinical
Phytoscience, 4, 32. Doi: 10.1186/s40816-018-0093-8
Pant, P., Pandey, S., & Dall’Acqua, S. (2021). The inuence of environmental
conditions on secondary metabolites in medicinal plants: A literature
review. Chemistry & Biodiversity, 18(11), e2100345. Doi: 10.1002/
cbdv.202100345
Pawar, R., & Rana, V. (2019). Manipulation of source-sink relationship in
pertinence to better fruit quality and yield in fruit crops: a review.
Agricultural Reviews, 40(3), 200-207. Doi: 10.3389/fenvs.2021.700768
Peñarrieta, J., Tejeda, L., Mollinedo, P., Vila, J., & Bravo, J. (2014). Phenolic
compounds in food. Revista Boliviana de Química, 31(2), 68-81. http://
www.scribd.com/bolivianjournalofchemistry
Pérez-Pérez, E., Castillo-Pirela, V., Ortega-Fernández, J., Sandoval-Sánchez, L.,
Medina-Lozano, D., Ramírez-Villalobos, M., & Ettiene-Rojas, G. (2020).
Catequina y epicatequina en hojas de guayabo Criolla Roja. Revista de
la Facultad de Agronomía de la Universidad del Zulia, 37(3), 262-279.
https://www.produccioncientificaluz.org/index.php/agronomia/article/
view/32662
Pérez-Pérez, E., Ettiene, G., Marín, M., Casassa-Padrón, A., Silva, N., Raga, J.,
González, C., Sandoval, L., & Medina, D. (2014). Determinación de
fenoles y avonoides totales en hojas de guayabo (Psidium guajava L.).
Revista de la Facultad de Agronomía de la Universidad del Zulia, 31(1),
60-77. https://www.produccioncienticaluz.org/index.php/agronomia/
article/view/27149
Pérez-Pérez, E. D. C., Ettiene-Rojas, G. R., Ramírez-Villalobos, M. D. C., &
Gómez-Degraves, Á. (2023). Sustancias antioxidantes en diferentes fases
fenológicas de Psidium guajava L. Agronomía Mesoamericana, 34(2),
52601-52601. Doi: 10.15517/am.v34i2.52601
Pérez-Pérez, E., Saavedra-Guillén, M., Ortega-Fernández, J., Sandoval-Sánchez,
L., Medina-Lozano, D., Ramírez-Villalobos, M., & Ettiene-Rojas, G.
(2019). Flavonoides en frutos de guayabo Criolla Roja (Psidium guajava
L.). Boletín del Centro de Investigaciones Biológicas, 53(3), 236-249.
Piasecka, A., Sawikowska, A., Kuczyńska, A., Ogrodowicz, P., Mikołajczak,
K., Krystkowiak, K., Gudyś, K., Guzy-Wróbelska, J., Krajewski, P., &
Kachlicki, P. (2017). Drought-related secondary metabolites of barley
(Hordeum vulgare L.) leaves and their metabolomic quantitative trait loci.
The Plant Journal, 89(5), 898-913. Doi: 10.1111/tpj.13430
Piñol, M., Palazón, J., & Cusidó, R. (2013). Introducción al metabolismo
secundario. En: J. Azcón-Bieto, M. Talón (Eds.), Fundamentos de
siología vegetal (pp. 323-348). (2 Ed.). McGraw Hill Interamericana.
Quirós, M., Petit, Y., Sánchez, A., Aponte, O., Poleo, N., Ortega, J., & Dorado, I.
(2009). Poblaciones de Oligonychus psidium Estebanes y Baker (Acari:
Tetranychidae) correlacionadas con aspectos fenológicos del guayabo
(Psidium guajava L.). Revista UDO Agrícola, 9(1), 208-216. https://
dialnet.unirioja.es/servlet/articulo?codigo=3293988
Rejeb, I.B., Pastor, V., & Mauch-Mani, B. (2014). Plant responses to simultaneous
biotic and abiotic stress: molecular mechanisms. Plants 3(4), 458-475.
Doi: 10.3390/plants3040458
Rioja Soto, T. C. (2020). Los metabolitos secundarios de las plantas y potencial
uso en el manejo de plagas agrícolas en agroecosistemas desérticos. Idesia
(Arica), 38(1), 3-5. https://revistas.uta.cl/pdf/349/1.pdf
Rivero-Maldonado, G., Pacheco, D., Martín, L., Sánchez-Urdaneta, A., Quirós,
M., Ortega, J., Colmenares, C., & Bracho, B. (2013). Flavonoides
presentes en especies de Psidium (Myrtaceae) de Venezuela. Revista de
la Facultad de Agronomía de la Universidad del Zulia, 30(2), 217-241.
https://www.produccioncientificaluz.org/index.php/agronomia/article/
view/27124
Salazar, D. M., Melgarejo, P., Martínez, R., Martínez, J. J., Hernández, F., &
Burguera, M. (2006). Phenological stages of the guava tree (Psidium
guajava L.). Scientia Horticulturae, 108(2), 157-161. Doi: 10.1016/j.
scienta.2006.01.022
Saltveit, M. E. (2017). Synthesis and metabolism of phenolic compounds. En:
E. M. Yahia (Ed.), Fruit and Vegetable Phytochemicals: Chemistry and
Human Health, 2nd Edition, (pp. 115-124). John Wiley & Sons, Ltd. Doi:
10.1002/9781119158042.ch5
Sampaio, B. L., Edrada-Ebel, R. A., & Batista Da Costa, F. (2016). Eect of the
environment on the secondary metabolic prole of Tithonia diversifolia:
A model for environmental metabolomics of plants. Scientic Reports, 6,
29265. Doi: 10.1038/srep29265
Shen, N., Wang, T., Gan, Q., Liu, S., Wang, L., & Jin, B. (2022). Plant avonoids:
Classication, distribution, biosynthesis, and antioxidant activity. Food
chemistry, 383, 132531. Doi: 10.1016/j.foodchem.2022.132531
Shivpoojan, A., Ram, R., Rajan, R., & Pandey, R. (2018). Ecacy of foliar
application of micronutrients on fruit set in winter season guava (Psidium
guajava L.) cv. Lalit. International Journal of Chemical Studies, 6(5),
2908-2910. https://dialnet.unirioja.es/servlet/articulo?codigo=8882545
Singh, S. (2011). Guajava (Psidium guajava L.). In: E. Yahia (Ed.), Postharvest
biology and technology of tropical and subtropical fruits, (pp. 213- 245).
Woodhead Publishing. Doi: 10.1533/9780857092885.213
Speed, M. P., Fenton, A., Jones, M. G., Ruxton, G. D., & Brockhurst, M. A.
(2015). Coevolution can explain defensive secondary metabolite diversity
in plants. New Phytologist, 208(4), 1251-1263. Doi: 10.1111/nph.13560
Tolić, M., Krbavčić, I., Vujević, P., Milinović, B., Jurčević, I., & Vahčić, N.
(2017). Eects of weather conditions on phenolic content and antioxidant
capacity in juice of chokeberries (Aronia melanocarpa L.). Polish Journal
of Food and Nutrition Sciences, 67(1), 67-74.
Tong, F., Medina, D., & Esparza, D. (1991). Variabilidad en poblaciones de
guayabo Psidium guajava L. del municipio Mara del estado Zulia. Revista
de la Facultad de Agronomía de la Universidad del Zulia, 8(1), 15-27.
https://www.produccioncientificaluz.org/index.php/agronomia/article/
view/25910
Turner, M. F., Heuberger, A. L., Kirkwood, J. S., Collins, C. C., Wolfrum, E.
J., Broeckling, C. D., Prenni J. E., & Jahn, C. E. (2016). Non-targeted
metabolomics in diverse sorghum breeding lines indicates primary
and secondary metabolite proles are associated with plant biomass
accumulation and photosynthesis. Frontiers in Plant Science, 7, 953. Doi:
10.3389/fpls.2016.00953
Verma, N., & Shukla, S. (2015). Impact of various factors responsible for
uctuation in plant secondary metabolites. Journal of Applied Research
on Medicinal and Aromatic Plants, 2(4), 105-113. Doi: 10.1016/j.
jarmap.2015.09.002
Vicente, O., & Boscaiu, M. (2018). Flavonoids: Antioxidant compounds for plant
defence…... and for a healthy human diet. Notulae Botanicae Horti
Agrobotanici Cluj-Napoca, 46(1), 14-21. Doi: 10.15835/nbha46110992
Vijaya Anand, A., Velayuthaprabhu, S., Rengarajan, R.L., Sampathkumar, P., &
Radhakrishnan, R. (2020). Bioactive Compounds of Guava (Psidium
guajava L.) In: H.N., Murthy, V.A. Bapat (Eds.), Bioactive Compounds
in Underutilized Fruits and Nuts (pp. 503–527). Springer International
Publishing. Doi: 10.1007/978-3-030-30182-8_37
Vranová, E., Coman, D., & Gruissem, W. (2012). Structure and dynamics of
the isoprenoid pathway network. Molecular plant, 5(2), 318-333. Doi:
10.1093/mp/sss015
Wang, L., Wu, Y., Bei, Q., Shi, K., & Wu, Z. (2017). Fingerprint proles of
avonoid compounds from dierent Psidium guajava leaves and their
antioxidant activities. Journal of Separation Science, 00, 1-13. Doi:
10.1002/jssc.201700477
Xu, H., Wang, G., Zhang, J., Zhang, M., Fu, M., Xiang, K., Zhang, M., & Chen,
X. (2022). Identication of phenolic compounds and active antifungal
ingredients of walnut in response to anthracnose (Colletotrichum
gloeosporioides). Postharvest Biology and Technology, 192, 112019. Doi:
10.1016/j.postharvbio.2022.112019
Yang, L., Wen, K. S., Ruan, X., Zhao, Y.Z., Wei, F., & Wang, Q. (2018). Response
of plant secondary metabolites to environmental factors. Molecules, 23,
762. Doi: 103390/molecules23040762
Zhang, Y., Butelli, E., & Martin, C. (2014). Engineering anthocyanin biosynthesis
in plants. Current Opinion in Plant Biology, 19, 81-90. Doi: 10.1016/j.
pbi.2014.05.011
