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A review on the influence of fertilizers application on grape yield and quality in the tropics

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Journal of Plant Nutrition
Authors:
  • Tanzania Agricultural Research Institute (TARI)
  • Tanzania Agricultural Research Institute, Makutupora Centre

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Grapevine nutrition has been a major focus for increasing grapes quality, productivity, and the way of minimizing the gap between actual and potential grape yield. Several nutrient elements are reported to influence grapevine physiological growth, yield, and grape quality. Being a perennial crop, the vines are fertilized with organic fertilizers to ensure a supply of organic carbon, improved soil organic matter content, and enhanced soil microorganisms. Globally, fertilizers with different NPK ratios and micronutrients are used to achieve a better yield and desirable quality of grapes. The deficiency of mineral nutrients results in chlorosis and eventual necrosis of leaves, which affect vine growth, yield, and grape quality in terms of sugar and polyphenols contents, consequently compromising wine quality and other grape-derived products. Thus, the establishment of the vineyard fertilizer program should be nutrient-specific, which takes into account vineyard management strategies that influence nutrient use efficiency from fallen leaves, pruning, soil testing, plant analysis, and other factors that affect yield and quality. However, applying mineral nutrients to grapevines may interfere with the physicochemical composition of grapes and derived products. Therefore, it is important to increase productivity, improve grape quality and ensure sustainable production through an integrated approach of using organic and inorganic fertilizers. This review covers the importance of organic fertilizer, macronutrients, and micronutrients applied to the grapevines for improving productivity and quality of grapes.
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A review on the influence of fertilizers application
on grape yield and quality in the tropics
Armachius James, Athuman Mahinda, Andekelile Mwamahonje, Elvillah
William Rweyemamu, Emmanuel Mrema, Kobusinge Aloys, Elirehema Swai,
Felista Joseph Mpore & Cornel Massawe
To cite this article: Armachius James, Athuman Mahinda, Andekelile Mwamahonje, Elvillah
William Rweyemamu, Emmanuel Mrema, Kobusinge Aloys, Elirehema Swai, Felista Joseph Mpore
& Cornel Massawe (2022): A review on the influence of fertilizers application on grape yield and
quality in the tropics, Journal of Plant Nutrition, DOI: 10.1080/01904167.2022.2160761
To link to this article: https://doi.org/10.1080/01904167.2022.2160761
Published online: 28 Dec 2022.
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A review on the influence of fertilizers application on grape
yield and quality in the tropics
Armachius James
a
, Athuman Mahinda
b
, Andekelile Mwamahonje
a
, Elvillah William
Rweyemamu
a
, Emmanuel Mrema
c
, Kobusinge Aloys
d
, Elirehema Swai
a
, Felista Joseph
Mpore
a
, and Cornel Massawe
a
a
Makutupora Center, Tanzania Agricultural Research Institute (TARI), Dodoma, Tanzania;
b
College of
Agriculture and Food Technology (CoAF), The University of Dar es Salaam, Dar es Salaam, Tanzania;
c
Tumbi
Center, Tanzania Agricultural Research Institute (TARI), Tabora, Tanzania;
d
Mlingano Center, Tanzania
Agricultural Research Institute (TARI), Tanga, Tanzania
ABSTRACT
Grapevine nutrition has been a major focus for increasing grapes quality,
productivity, and the way of minimizing the gap between actual and
potential grape yield. Several nutrient elements are reported to influence
grapevine physiological growth, yield, and grape quality. Being a perennial
crop, the vines are fertilized with organic fertilizers to ensure a supply of
organic carbon, improved soil organic matter content, and enhanced soil
microorganisms. Globally, fertilizers with different NPK ratios and micronu-
trients are used to achieve a better yield and desirable quality of grapes.
The deficiency of mineral nutrients results in chlorosis and eventual necro-
sis of leaves, which affect vine growth, yield, and grape quality in terms of
sugar and polyphenols contents, consequently compromising wine quality
and other grape-derived products. Thus, the establishment of the vineyard
fertilizer program should be nutrient-specific, which takes into account
vineyard management strategies that influence nutrient use efficiency from
fallen leaves, pruning, soil testing, plant analysis, and other factors that
affect yield and quality. However, applying mineral nutrients to grapevines
may interfere with the physicochemical composition of grapes and derived
products. Therefore, it is important to increase productivity, improve grape
quality and ensure sustainable production through an integrated approach
of using organic and inorganic fertilizers. This review covers the import-
ance of organic fertilizer, macronutrients, and micronutrients applied to
the grapevines for improving productivity and quality of grapes.
ARTICLE HISTORY
Received 1 August 2022
Accepted 7 November 2022
KEYWORDS
fertilizers; Grapes; grapevine
nutrition; Vineyards;
Viticulture
Introduction
Grapevine (Vitis vinifera L.) is one of the most economically important productive drought
stress-adapted crops grown globally (Gambetta et al. 2020). Grapes are considered one of the
most popular and favorite fruits in the world for their excellent flavor, taste, and nutritional
value. They are mainly consumed as fresh fruits, juice, raisins, and wine. Grapes consist of sugar
in form of monosaccharides (glucose and fructose) and various phenolic compounds (Kandylis,
Dimitrellou, and Moschakis 2021; Li et al. 2020). It also contains minerals such as potassium,
magnesium, iron, and calcium as well as vitamin C, B12, and B6, which are nutritionally import-
ant (Conde et al. 2007; Pezzuto 2008). Achieving a balance between productivity and fruit quality
CONTACT Armachius James armachiuss@gmail.com Makutupora Center, Tanzania Agricultural Research Institute (TARI),
Dodoma, Tanzania
ß2022 Taylor & Francis Group, LLC
JOURNAL OF PLANT NUTRITION
https://doi.org/10.1080/01904167.2022.2160761
is a major goal in viticulture, which is particularly a challenge considering the threats of climate
change and weather variability. Among other factors; soil nutrient availability plays a major role
in grapes productivity and quality, which creates the need for the use of fertilizers. The most
important nutrients for the vines are nitrogen (N), potassium (K) and phosphorus (P). Other ele-
ments include Carbon (C), Chlorine (Cl), sulfur (S), magnesium (Mg), boron (B), zinc (Zn),
manganese (Mn), copper (Cu), iron (Fe), silicon (Si), calcium (Ca), and molybdenum (Mo),
which play the role in grapevine physiological growth, yield, and quality (Brunetto et al. 2015;
Karagiannidis et al. 2007; White and Brown 2010). Their subsequent effects on must fermentation
and wine quality have been identified, giving even greater importance to vineyard nutri-
tional management.
Nutrients occur naturally in the soil and are taken up by the plant roots. Too high level of
nutrients causes vigorous growth of the vine producing a dense canopy that excessively shades
the fruit or shows a varied range of toxicity. On the other hand, the soil nutrients are depleted
overtime, and need to be replenished with organic and/or mineral fertilizers to improve grapes
yield and quality. Consequently, the common symptom linked to lack of nutrients is chlorosis,
such that the color of leaves changes from yellow to purple, hence restricted photosynthesis and
grapes struggle to ripen properly, while quality and yield are reduced. Moreover, inadequate soil
nutrients may lead to increased susceptibility of the grapevines to pests and/or diseases, which
demand a large quantity of pesticides to be applied to manage them. The problem can be
addressed by using appropriate fertilizers, particularly a mixed formulation that provides macro
and micronutrients (Gur et al. 2022). The most immediate effect, which can raise the yield and
wine quality are crop load and NPK levels (Fiaz et al. 2021; Michopoulos and Solomou 2019;
Solomou, Michopoulos, and Vavoulidou 2022). Interestingly, berry N and to some extent P and
K concentrations can be manipulated, intentionally or otherwise by viticultural and oenological
factors. Thus, grapes characteristics are controlled by cultivation practices including weeding, pest
control, crop load, irrigation, canopy management, and fertilizer application. Moreover,
Hasanaliyeva et al. (2021), reported grape variety as the strong explanatory variable for total phe-
nols and anthocyanins.
The application of organic fertilizers represents the standard traditional method for vineyard
fertilization, which is characterized by the gradual release of the bound nutrient elements during
the mineralization process contrary to inorganic fertilizers. Organic fertilizers contribute signifi-
cantly to preserving natural resources, increasing soil organic carbon, improving soil structure,
minimizing the adverse effects of mineral fertilizers on the environment, and sustaining other
organisms, which are environmentally friendly for sustainable agroecology (Ball et al. 2020;
Lazcano et al. 2022; Wilson, Lambert, and Dahlgren 2021). Thus, organic fertilizers prove further
benefit in enriching the soil microbes, a balanced pH, and a range of minerals. Despite the bene-
ficial effect on soil structure, manure has no significant effect on crop production compared to
chemical fertilizers (Mahinda et al. 2018). For instance, farmers in Tanzania have traditionally
used only organic manure (farmyard manure and compost), although the combined application
of mineral fertilizers has been recommended in most vineyards. Organic and mineral fertilizers
are important for the production of grapes and attaining yield and quality, which complement
each other in maintaining soil fertility and sustaining high yield. The integrated fertilization
approach provides a transition between conventional and organic fertilization, while maintaining
yield and sustainably decreasing the residues of mineral fertilizers in the environment. For
instance, Michopoulos and Solomou (2019), recommended foliar application of Mg and Zn for
the organic vineyards to attain nutrient sufficiency. PoPopovi
c et al. (2020), reported improved
grapes yield, physical and chemical characteristics of bunches and berries following application
of 50% organic and 50% mineral fertilizers (NPK 8:16:24) on the Cardinal grape vineyard.
Similarly, Chowaniak et al. (2021), reported increased yield on application of N fertilizers as a
single dose or divided into three consecutive doses. Also, Koureh et al. (2019), reported an
2 A. JAMES ET AL.
increased yield of grapes following application of fertilizers, although organic fertilization systems
resulted in grapes with the highest nutritional quality, antioxidants, total flavonoids, and valuable
phenolics. On the other hand, excessive mineral fertilizer application, especially nitrogen fertil-
izers, may lead to the accumulation of harmful residual substances such as nitrites in the grape
leaves and berries. The long-term use and/or uncontrolled application of inorganic fertilizers can
cause soil and water pollution, leading to the deterioration of the soil structure and reduction of
soil microflora. The study of Wu et al. (2021), reported reduced fungal diversity on the grape
berry surfaces from the vineyard fertilized with mineral fertilizers in the study conducted in
Xinjiang, China.
Overall, there is limited information about the impact of inorganic fertilizers on grape prod-
uctivity and quality, particularly in the tropics as compared to other crops. Therefore, this study
aimed to review the use of inorganic fertilizers to improve the productivity and quality of grapes
as adequate nutrition is essential for the growth and yield of the grapevines.
Organic fertilizer for vineyard nutrition
Organic fertilizers are fertilizers derived from organic sources including animal or plant based
materials that are either by-products or end products of naturally occurring processes such as
manure, plant residues, and composted organic materials (Longa et al. 2017; Michopoulos and
Solomou 2019). Organic fertilizers are essential source of plant nutrition as they gradually release
plant nutrients through the mineralization process, which maintains a balance for healthy growth
of grapevines while enhancing healthy soil. They reduce necessity of repeated application of syn-
thetic fertilizers to maintain soil fertility so as mitigating problems associated with synthetic fertil-
izers (Mahinda et al. 2018). The commonly used organic fertilizers for grape production include
farmyard manure, compost, vermicompost, sewage sludge, food processing wastes, and biosolids.
In the tropics, and Tanzania in particular, farmyard manure has been a choice for vineyard
fertilization during initial trench filling, planting and for amendment applications. Likewise, com-
post maybe opted during initial vineyard management. Application of organic fertilizers help to
increase soil water holding capacity, improve soil texture, organic matter content, and a nutrient-
rich ecosystem (Longa et al. 2017; Wilson, Lambert, and Dahlgren 2021). Farmyard manure
mainly consists of decomposed mixture of animal excreta (urine and dungs) mostly from cow-
dung, bedding materials, soil, and sometimes dropped feeds depending on specific livestock man-
agement practices (Khoshnevisan et al. 2021; Rayne and Aula 2020). Organic fertilizers are
greener option for plant nutrition as play a pivot role in the protection and improvement of the
soil environment and promote growth of soil microorganisms (Semenov et al. 2021). Moreover,
farmyard manure provides a buffering capacity for the soil pH that control microbial community
composition and mediate nutrient availability for plant growth in the vineyard (Liu et al. 2020).
Given the continued development of modern agriculture, the role of organic fertilizers in viti-
culture is becoming more important and an integrated part for sustainable grape production
(Balafoutis et al. 2017; Lamastra et al. 2016). The use of farmyard manure integrated with deep
trenching provides an effective way to avoid and combat soil compaction in the vineyard (van
Antwerpen et al. 2021). Specifically, grapes harvested from the vineyard applied with organic fer-
tilizers have good taste and can effectively maintain the unique nutrition and flavor as well as a
characteristic wine terroir (Lazcano, Decock, and Wilson 2020; Liu et al. 2019). However, if
manure is over-applied and not well incorporated might be carried away with runoff to water
bodies affecting water quality and aquatic systems. Conversely, manure can emit greenhouse gases
such as methane (CH
4
) and nitrous oxide (N
2
O), which are known to significantly contribute to
climate change events (Skinner, Gattinger, et al. 2019). Hence proper management, avoidance of
over-application and the use of controlled release of organic fertilizers are effective and advanced
JOURNAL OF PLANT NUTRITION 3
ways for a sustainable vineyard. Granted that, there is a need for continued research and develop-
ment on utilization and management of organic fertilizers.
Vineyard nutrition and fertilizer management
Managing the nutritional requirements of the vineyard requires a visual assessment of the vines,
growth habits, and the nutrient status of plant tissues and/or soil to develop an appropriate fertil-
izer program (Arrobas et al. 2014). Each vineyard may have a unique combination of soil type,
vine age, canopy architecture, and cultivar, thus nutritional requirements vary among vineyards,
even locations within a vineyard. Analysis of the nutrient content of the petioles gives a good
indication of available nutrients to the plant. The petiole nutrients are compared to a standard
representing deficient, adequate or high levels of specific elements on plant performance (Zhao
et al. 2019). Whereas, soil analysis gives nutrient composition, which may not necessarily be
available for uptake by the grapevine. Additionally, soil physical properties such as soil texture
and structure may also be assessed as they influence nutrient availability. Collectively, sandy soils
are likely to be leached of nutrients, and high clay content soils will rapidly fix applied potassium
fertilizers. The soil high in organic content has high levels of readily available nutrients.
Evidently, high microbial biomass, organic carbon, exchangeable K, Ca and Mg, increased P and
N availability, a balanced pH, and improved yield were reported in the vineyard treated with
compost (Ball et al. 2020; Wilson, Lambert, and Dahlgren 2021).
Vineyard nutrient deficiency can be improved by supplementation. However, high nutrient
concentration can be toxic and produce an adverse effect on grapevine performance, grape quality
and their derived products (Serrano et al. 2017). Grapevines require the supplementary applica-
tion of fertilizer to ensure maximum production and sustainable grape quality. The use of inor-
ganic fertilizers provides better results because of the application of a known and exact amounts
of fertilizers to the grapevine. In absence of limiting factors like water, the application of inor-
ganic fertilizers provides an easily available source of micronutrients for the vine. Macronutrients
are usually applied to the soil surfaces in the dry form, ripped into the soil or via fertigation. The
most important macronutrients are N, P and K, which directly affect plant growth and create
various plant parts. These nutrients, in particular N and K have high mobility and can be translo-
cated rapidly from older to new leaves if the demand in young leaves or fruiting bodies is higher
than the supply from the soil, resulting in chlorosis and necrosis (Jegadeeswari, Chitdeshwari,
and Shukla 2020; Michopoulos and Solomou 2019). However, the case is not the same for micro-
nutrients as they are generally applied directly to the vegetative part via foliar sprays, and are
required in trace amounts Table 1. Generally, these nutrients interact with each other to effect
several pathways in the plant (Kumar, Kumar, and Mohapatra 2021). A successful fertilizer man-
agement program depends on the selection of the correct fertilizer to address the specific deficit
observed in the vineyard, the correct rate of nutrient requirement and the timing for application
Table 1. List of essential nutrients for the grapevines.
Macronutrients Micronutrients
Carbon (C) Iron (Fe)
Hydrogen (H) Zinc (Zn)
Oxygen (O) Boron (B)
Nitrogen (N) Manganese (Mn)
Phosphorus (P) Copper (Cu)
Potassium (K) Molybdenum (Mo)
Calcium (Ca) Chlorine (Cl)
Magnesium (Mg) Nickel
Sulfur (S) Cobalt (Co)
Silicon (Si)
Sodium (Na)
4 A. JAMES ET AL.
of the fertilizer (Rahaman et al. 2019). Collectively, providing several benefits in terms of soil and
environment protection, while enhancing the grape quality and yield.
Timing of fertilizer application
The timing of fertilizer application depends on the vine age, phenology, soil moisture, nutrient
mobility and cation exchange capacity. The young grapevines require increased nitrogen for rapid
root and vegetative growth compared to older and established vines. The phenological develop-
ment of a grapevine dictates the types and amount of nutrients required based on the growth of
vegetative or reproductive components of the vines (Cameron, Petrie, and Barlow 2022; Parker
et al. 2014; Stefanello et al. 2021). Either during budding and budburst, bloom, fruit set, v
eraison,
maturity to leafy senescence or subsequent leaf fall (Neilsen et al. 2010). For instance, N, P, K,
and Mo are important after budburst, when the vines are undergoing rapid vegetative growth.
Zinc, Mn, Mg, Fe, and B are critical to flowering when compounds are required for fruit set in
the current season and good bud initiation for the following season (Sabir et al. 2014; Usha and
Singh 2002). After harvest, root growth and carbohydrate storage in the trunk may require the
addition of N, P, and Ca.
Mode of fertilizers application on the grapevine
The mode of fertilizers application determines fertilizer use and recovery efficiency as well as dis-
tribution in the grapevines. Chemical fertilizers can be applied to plants by broadcasting, banding,
spot, fertigation and/or foliar application (Likar et al. 2015; Romic et al. 2012; Topalovi
c et al.
2011). The study by Williams (2015), applied N fertilizers in the form of potassium nitrate
(KNO
3
) or ammonium sulfate (NH
4
)
2
SO
4
in the drip or fallow irrigated grapevines shortly after
anthesis with a single application of 25 g N/vine (28 kg N/ha) or split application with labeled
KNO
3
(31 kg N/ha) during the growing season. Williams (2015), found that approximately 47%
and 30% recovery of the (NH
4
)
2
SO
4
and KNO
3
fertilizer, respectively in the drip-irrigated clusters
and only 12% in the fallow irrigated clusters indicating fertilizer type and mode of application
determines the fertilizer recovery efficiency and the distribution of N in the grapevines. At the
same time, foliar application of
15
N-labeled urea (20 kgN/ha) has shown effective fertilizer utiliza-
tion and recovery in the grapevine, while maintaining grape quality and increased must yeast
assimilable nitrogen (YAN) and K (Verdenal et al. 2022).
Fertigation with drip-irrigated systems
In drip-irrigated vineyards, fertigation is generally the most effective way to apply N, P, and K as
it allows fertilizers direct to the rhizosphere and offers the economical use of water (Gong,
Zhang, and Liu 2022; Verdenal et al. 2022). Because the nutrients are placed into moist soil with
active roots, K or P uses efficiency generally is increased as compared to a band application in a
furrow close to the grapevines (Christensen, Peacock, and Bianchi 1991; Davenport and Jones
2016; Schreiner and Osborne 2018). In the vineyards with a drip-irrigated system, K can be ferti-
gated during the growing season, with the application being made during bloom or before
v
eraison. However, it may be more practical to apply K every week than to apply a large amount
with one single application. Other nutrients including Ca, Mg, Zn, Cu, Fe, and Mo can also
be fertigated.
Furrow irrigated systems
Under the furrow irrigated system, fertilizer is placed near the outer edges of the furrow toward
the grapevine rows. Nitrogenous fertilizers such as urea and ammonium forms, should always be
JOURNAL OF PLANT NUTRITION 5
drilled at least two inches deep into the soil or immediately incorporated, since they are subjected
to volatilization losses if left on the surface (Stefanello et al. 2021). For young vines planted on
sandy soil, fertilizer should be applied within three feet of the row to ensure root access.
Nevertheless, in K deficient sandy soils, grapevines respond more rapidly to surface applied K
compared to furrow application. This may have been because, during the first year after surface
application, most K is confined to the top foot of the profile where the root density is high.
Potassium demand is highest between bloom and v
eraison, during this period 60% of the annual
K demand is taken up (Baby et al. 2022; Ciotta et al. 2016). Between v
eraison and harvest, only
about 10% of the annual K demand is taken up. This period is characterized by the redistribution
of K from vegetative organs to clusters (Baby et al. 2022). About 15% of the annual K demand
may be covered with postharvest uptake.
Foliar application
Foliar nutrient application to grapevines gives advantages for economical and environmental rea-
sons. It can improve nutrient uptake efficiency as nutrients are directly absorbed by leaves. Foliar
spraying provides specific nutrients to the grapevine on a timely basis during the critical stage of
growth like budding, flowering and fruiting (Gautier et al. 2018; Masi and Boselli 2011;P
erez-
Alvarez et al. 2017). A well-planed grapevine foliar nutrition program can supplement soil fertil-
izers especially when the vine root system is unable to keep up with crop demand or when soil
nutrients are unavailable. For instance, fertigation, furrow or broadcasting are subjected to loss to
the environment via leaching, denitrification, surface runoff, gaseous emission and microbial con-
sumption. Previous studies reported that foliar N did not increase leaf N concentration, vigor,
TSS nor yield in the vineyard with adequate soil N. In that regard, foliar N cannot replace soil or
irrigated water applied N because the quantity of N that can be taken up by the leaves is too little
to cover the seasonal requirements. Yet, foliar N application may provide some benefit when
plants are N deficient during the growing season depending on the severity of N deficiency. Kelly
et al. (2017), reported increased berry YAN and amino acids following foliar N and S at a rate of
15 and 5 kg/ha, respectively applied in two splits before v
eraison.
Foliar N application during v
eraison could prevent nitrogen deficiency in grapes and must
from N deficient vineyards. Also, foliar N can increase YAN and phenolics and improves certain
flavor components of grapes and wine (Cheng et al. 2021). Previous studies found that foliar N
performs better at the beginning of v
eraison and one or two weeks later; the reason being, that
foliar N provides nutrients to the most vigorous part (the grape berries) of reproductive growth
metabolism. At this stage, vegetative growth slows or even stops, and grapevine growth concen-
trates on the grape berries. To some extent, the leaf surface during v
eraison is old enough that
some cracks may exist on the leaf wax surface, resulting in the promotion of nutrient uptake.
Thus, a small amount of leaf N supplement can promote the accumulation of nitrogen in the ber-
ries and meet N needs for the grape in a timely and efficient manner. At the same time, moderate
N supply during v
eraison may increase phenylalanine content, which is the precursor of phenolics
and contributes to the accumulation of anthocyanins, flavanols, and stilbenes. During v
eraison,
essential growth is focused on the berry development, so supplemental N may lead to an increase
in photosynthesis of grapevine and promote storage of excess photosynthesis products mainly
carbohydrates in the berries, benefiting the synthesis of secondary metabolites (Cheng et al. 2021;
Stefanello et al. 2020,2021). From the v
eraison to ripening ammonium (NH
4
þ
) content in the
grape berries gradually decreases, thus foliar N application may increase NH
4
þ
in grapes.
On the other hand, the quantity of K required by grapevines is too large to be supplied
through the leaves. Similarly, the benefits of foliar K application are minimal. Other studies did
not find effects of foliar K applications on grapevines growth, yield or fruit quality. Moreover,
most of the micronutrients have shown effective results through foliar application methods as
detailed in section five.
6 A. JAMES ET AL.
Macronutrient fertilizers
Nitrogen
Nitrogen (N) is an essential soil-derived macronutrient element for plant growth. It is taken up
by plants as nitrate. Nitrogen, among nutrients that soil supplies to grapevines, is the most influ-
ential on grapevine physiological growth, vigor, yield as well as grape and must quality. It is a
part of proteins and nucleic acids. Nitrogen is involved in the winemaking process from grape-
vine growth to wine fermentation, and its precise utilization in the vineyard can regulate grape
and wine quality (Arrobas et al. 2014). During winemaking, N in the must can nourish yeast
growth and metabolism and ultimately modify the flavor components in wine, which influences
the quality and sensory attribute of wine. It supplies nitrogen in form of amino acids and yeast
YAN for yeast growth and fermentation performance (Gong, Zhang, and Liu 2022; Holzapfel
et al. 2015; Miliordos et al. 2022). Nitrogen deficiency may cause several disorders leading to
overall reduced plant performance. Though, grapes have a low-N fertilizer requirement of 84 kg
N/ha compared to most other crops of approximately 125-275 kg N/ha (Tassinari et al. 2022;
Williams 2015). The lower requirement of N fertilizers can be explained that N may be supplied
from the reserves in the permanent plant structures of the grapevines. For instance, Brunetto
et al. (2016), revealed that by applying urea at the budding and full bloom stage, grapevines took
up more N; when the application rate was 20 kg N/ha during budding in the first season,
although most N contained in the plant organs was derived from other sources than the fertilizer
N. The N-derived from fertilizer applied at different rates was recovered preferentially in the
leaves and fruits in the following season. Similarly, Gong, Zhang, and Liu (2022), revealed that
the absorption of fertilizer N by grapevine accounted for 12-17% of the total N absorption, and
the proportion from the soil N was as high as 82-87%.
The dynamics of N uptake and the impact on the physiology and grapevine yield are necessary
for the proper application of nitrogenous fertilizers. Proper application of N fertilizers ensures
not only high yield but also, balanced vegetative and reproductive growth in the plants. The
increase of grape cluster weight and length as well as yield depends on the N doses (Stefanello
et al. 2020). For instance, the application of N fertilizers over 20 kg N/ha, results in the highest
yield and higher phenolics composition in Cabernet Sauvignon berries (Lorensini et al. 2015).
While, Stefanello et al. (2020), reported decreased anthocyanin concentration in the must from
grapes fertilized with N fertilizers, which can be explained as dilution effect on the pulp/skin
ratio, promoted by the increased yield. Delaying N application until the bloom is associated with
decreased canopy density and increased yield. The application of N during bloom for Merlot or
budburst for Chardonnay and Pinot Noir has been a strategy for targeting berry N status without
causing excess vegetative growth (Tassinari et al. 2022). Likewise, increasing N supply, increases
plant vigor, chlorophyll and stomata aperture because N stimulates photosynthesis. Generally, N
is the most potent in its ability to influence grapevine physiological growth, morphology, fruit
production, and tissue composition as it forms part of several cell constituents including proteins,
chlorophyll, and nucleic acids (Arrobas et al. 2014). Thus, increased soil N availability enhances
photosynthesis, which means more sugar for growth and fruit ripening.
Imbalanced use or widespread use of N fertilizers alone favors vegetative growth of lateral
shoots hence dense canopy associated with reduced fruit sugar and delayed ripening, which
causes poor quality of the fruits in terms of total soluble solids (TSS), color, keeping quality, min-
eral content, high malic acid, low must pH, and the fruits become susceptible to sunburn (Cocco
et al. 2021; Conradie and Saayman 1989). High soil N, inhibit root growth, which makes grape-
vines more susceptible to water stress, pest and diseases. Thus, nitrogenous fertilizers should be
applied to ensure an adequate N supply during spring development, but available N in late sum-
mer should not be high enough to encourage late season shoot growth, delayed maturing and
promote immature canes.
JOURNAL OF PLANT NUTRITION 7
Excess nitrogen in grapevine
Excess N is characterized by dark green leaves, reduced fruit set, and delayed ripening of grapes.
High N availability results in high canopy vigor with excessively long shoots. The increased can-
opy density in turn reduces light interception in the fruit zone, which generally results in delayed
fruit ripening and reduced concentration of total soluble solids (TSS), anthocyanins, terpenes, and
total phenols in grapes (Schreiner, Scagel, and Lee 2014). High N uptake can also increase the
susceptibility of the grapevines to diseases, such as powdery and downy mildew, and bunch stem
necrosis in which developing flower clusters are aborted by the vine causing severe yield reduc-
tion. This is in alignment with the study of Add El-Razek et al. (2011), who revealed that high N
results in increased juice per berry and decreased fruit firmness. However, the relationship
between N status and the berry composition remains complex as depends on many other factors.
For instance, an interaction between N and K fertilizers was previously presented; a moderate K
and increased rate of N supply decreased grape skin polyphenols, while increasing N at a high
rate of K supply increased berry skin polyphenols.
Nitrogen deficiencies symptoms in grapevine
Nitrogen deficiency results in reduced vegetative growth and photosynthesis, small-thin and stiff
leaves, pale-green to yellow leaves, slowed shoot growth, shortened internodes, and low berry set
(Cocco et al. 2021; Verdenal et al. 2021). Consequently, the berries mature before they reach their
normal size packed with low sugar content and thus reduced yield (Schreiner, Lee, and Skinkis
2013). Nevertheless, N deficiency enhances phenolics and tannins in the berries (Schreiner,
Scagel, and Lee 2014). Leaf symptoms appear after the beginning of ripening as the result of N
translocation from the leaves to the berries Figure 1. When N deficiency is accompanied by water
stress, the leaf margins may roll slightly upwards, wilt, and dry. Sometimes tissue between the
veins turns light brown and dies. Under extreme N deficiency, the leaves wilt and drop.
During fermentation, N plays an important role as a food source for yeast growth and per-
formance. Therefore, low grape nitrogen content impact the fermentation rate such that lessened
YAN content in the must and inhibits yeast growth and metabolism, which could lead to stuck
and sluggish fermentations (Paul Schreiner, Osborne, and Skinkis 2018; Tian et al. 2022). For
instance, must with less than 140mgN/l of YAN is prone to stuck and sluggish fermentation asso-
ciated with low sensory quality and flavor problems such as rotten egg hydrogen sulfide (H
2
S),
branched fatty acid ethyl esters, volatile mercaptans as well as increased alcohol (Cheng et al.
2021). Hydrogen sulfide is generated from the breakdown of protein by the starving yeast cells.
The addition of diammonium phosphate (DAP) is a common practice for increasing must N lev-
els and maintaining moderate fermentation vigor (Tian et al. 2022). On the other hand, high N
may not necessarily benefit wine quality as may lead to the accumulation of ethyl carbonate and
biogenic amines and encourage microbial spoilage and instability.
Figure 1. Leaves with nitrogen deficiency symptoms.
8 A. JAMES ET AL.
Types of N fertilizers for grapevines
Different nitrogenous fertilizers with their percent nitrogen content include urea (46% N), calcium
ammonium nitrate (CAN 26% N), potassium nitrate, ammonium sulfate (SA), ammonium sulfate
nitrate (ASN (27% N)), diammonium phosphate (DAP), and ammonium bicarbonate (ABC) are
applied to plants to supply N (Zhang et al. 2011). Urea and ammonium fertilizers, especially ammo-
nium sulfate acidify soils over time and should be avoided on soil with a pH below 6. At the same
time, urea promotes rapid volatilization while nitrate fertilizers contribute to leaching (Cao, Lu, and
Yu 2018). Grapevines fertilized in the fall should have an active, healthy canopy for three to four
weeks after application. When uptake time is limited, a nitrate fertilizer, which is mobile and imme-
diately available, may be the best choice. When the canopy is no longer active, N is not taken up
and may be leached below the root zone during winter. The uptake of N is relatively slow between
budbreak and bloom, but high between bloom and v
eraison. During this period, about half of the
annual N demand is taken up. Due to the ability of grapes to mobilize reserves from permanent
structures, time of application has less effect on grapevine performance than it has in annual crops,
provided the N is applied during a period of active uptake. Studies have shown that, N is best
applied in spring during a period starting after budbreak until fruit set. During flowering to
v
eraison, the vine accumulates most of the N as is associated with cell division and growth, hence
required for the synthesis of chlorophyll, nucleic acids and proteins, and achieve the highest vine
mass (Ferrara et al. 2018). In the temperate climate, early application before or at budbreak is sus-
ceptible to leaching from sprinklers for frost protection, late rainfall or excessive spring irrigation
should be delayed until after the frost danger period. Nitrogen applied between harvest and leaf
senescence refills storage reserves in permanent structures and supports leaf growth the following
spring, in the regions where the leaf canopy remains active for an extended period after harvest.
Nitrogen application in late fall after leaf falls is inefficient, because N may be leached below the
root zones by rains. Therefore, studies on the amount, duration, and methods of N application on
the grapevine are imperative for yield and quality improvement.
Phosphorus (P) (P
2
O
5
,H
2
PO
4-
) and phosphate fertilizers in grapevines
Phosphorus (P) available as anion phosphate (H
2
PO
4-
) is required for energy metabolism (ATP)
and nucleic acid synthesis: the deoxyribonucleic acid (DNA) and ribonucleic acid (RNA)
(Davenport and Jones 2016). The most commonly used form of phosphorus fertilizers includes
diammonium phosphate (DAP), monoammonium phosphate (MAP), and monocalcium phos-
phate (MCP) (Meyer et al. 2020). It plays a phosphorylation role in photosynthesis, respiration
and regulation of enzymes. Phosphorus is involved in the formation of cellular membranes such
as plasma and the vacuole membrane. It plays a vital role in stimulating root development par-
ticularly seminal and hair roots formation that increases water and nutrient absorption as well as
an increased stalk, stem length, and resistance to plant diseases (Schreiner and Osborne 2018;
Zheng et al. 2020). It maintains quality grapevine flowering and fruiting. However, the P require-
ment for a grapevine is generally low.
Application of P fertilizers on the P deficient vineyard of Cabernet-Sauvignon, Chardonnay
and Chenin Blanc resulted in increased leaves and juice P (Skinner, Gattinger, et al. 2019). The
application of P fertilizers on grapevines was also reported to influence differences in appearance,
flavor, aroma, and taste in wine made from Sauvignon, Chardonnay, and Chenin Blanc. At the
same time, increased P status in vine increases wine anthocyanins and soluble phenolics and fer-
mentation rate in the Cabernet-Sauvignon (Skinner, Gattinger, et al. 2019).
Phosphorus deficiencies in grapevine
Deficiency of P leads to stunted plant growth, weakened roots, reduced shoot growth, darkened
purple-to-red leaves and yellowing of the interveinal area of basal leaves. Under severe P
JOURNAL OF PLANT NUTRITION 9
deficiency, young leaves may turn reddish, whereas red spots near the edges may appear on older
leaves (Schreiner and Osborne 2018). While veins remain green, the area between them turns
red, appearing as islands or bars of red tissues Figure 2. Phosphorus deficiency reduces fruit set,
resulting in loose and small clusters (Davenport and Jones 2016). P availability may be limiting in
the vineyard at higher elevations on marginal, shallow, and acidic soils. Notably, P deficient
grapevines have been found in vineyards located on hillsides.
Potassium fertilizers (K) in grapevine
Potassium (K) is an essential element for plant nutrition and its ability to influence meristem
growth, water uptake, photosynthesis, and long-distance transport of assimilates via xylem and
phloem (Wang and Wu 2015). Potassium forms a major osmotic solute of the grapevines. Owing
to its fundamental roles in turgor generation, primary metabolism, and long-distance transport, K
plays a prominent role in crop resistance to drought, salinity, high light or cold as well as resist-
ance to pest and fungal diseases (Rogiers et al. 2017). Potassium accumulates primarily in the
berry skin tissues during ripening as a result of K remobilization from mature leaves. Grape ber-
ries are a strong sink for K, particularly during ripening. The most abundant cation in grape ber-
ries is K, which contributes to charge balance and may be involved in sugar transport. Potassium
reduces acid levels in berries and interacts with tartaric acid to form potassium bitartrate, which
has limited solubility. A high potassium supply increases the TSS and decreases the total acidity
(TA) of the berries (Ben Yahmed and Ben Mimoun 2019; Ciotta et al. 2016). Adequate potassium
nutrition helps to increase both the coloring and polyphenol content of berries. Other beneficial
effects of K include high juice content, high vitamin C content, uniformity and acceleration of
ripening of fruits and resistance to bruising or physical damage during shipping and storage
(Griesser et al. 2017). Several research results suggest the role of K in enhancing the quality of
fruits and their keeping, and marketable quality.
Applying K fertilizers improves yield and grape quality, vitamin C content, and increased total
sugar slightly, while the acidity is reduced. Increasing N-K rates increase pH and decrease TA.
This mechanism seems to be that excessive K migrates to the fruit and enhances the formation of
potassium bitartrate, which precipitates and causes a decrease in TA. Generally, oversupply of K
resulting from excessive fertilizer use can lead to high berry K concentration and quite often high
must pH.
K Deficiencies in grapevine
In the K-deficient vineyards, the supply of sink organs with photosynthates is impaired and sugar
accumulates in source leaves. Limits shoot and fruit growth and results in premature leaf fall,
poor fruit set, decreases fruitfulness, delayed ripening, and low yield (Griesser et al. 2017; Rogiers
et al. 2017). Also, low K supply results in low fruit K concentration as well as low must pH
Figure 2. Leaves with phosphorus (P) deficiency symptoms.
10 A. JAMES ET AL.
(Schreiner, Lee, and Skinkis 2013). Chlorosis may occur briefly at the tip and margin of older
leaves as well as necrotic spotting across old leaves blades Figure 3. Overall, the vine becomes sus-
ceptible to powdery mildew. However, the impacts of mild K in the grapevine, show no visual
differences in leaves or fruit clusters.
Types of K fertilizers
Various K fertilizers can be used in vineyards, with potassium sulfate (K
2
SO
4
) being the most
popular. Others include potassium chloride (KCl), potassium nitrate (KNO
3
), and potassium
magnesium sulfate (Rawat, Sanwal, and Saxena 2016; Teotia et al. 2016). Drainage problems or
high soil chloride concentrations may prevent the use of KCl in some areas. In general, chloride
fertilizers should be avoided in areas with salinity concerns. Potassium carbonate has an alkaline
soil reaction and is often used in acidic soils to increase soil pH. Potassium thiosulfate, on the
other hand, acidifies the soil and is used in alkaline soils Table 2.
Other macronutrients for grapevine productivity and grape quality
Magnesium (Mg), Calcium (Ca) and Sulfur (S) are also found in the grapevine tissues and con-
tribute to the functioning and structure of the vine.
Magnesium (Mg)
Magnesium (Mg) is an important macronutrient with several physiological functions that influ-
ence plant physiological growth and development. It is the central atom of chlorophyll and it acti-
vates enzymatic processes, thus contributing to carbohydrate production in leaves through
photosynthesis. Also, favorably influences assimilation, such that, high Mg levels may limit the
uptake of K by the vein. The study by Zl
amalov
a et al. (2016), emphasized that foliar application
of Mg as magnesium sulfate (MgSO4) at 3.86 kg Mg/ha increased grape yield by 11.2% compared
to control while other parameters were insignificant. Magnesium uptake by the plant is also
affected by the antagonistic effect of Ca and K and vice versa. For instance, three times applica-
tions of foliar Mg totaling 2.5 kg Mg/ha on the grapevine grown on the calcareous soil, revealed
that Ca is dominant in the plant disturbing K and Mg physiology (Gluhi
c et al. 2009).
Magnesium deficiency reduces chlorophyll content in the leaves and changes the chlorophyll
a:b ratio in favor of chlorophyll b. Magnesium symptoms appear in mid- to late-season and
include marginal leaf yellowing, reddening of basal leaves and extending to the internal vein area,
while the mid-vein region remains green Figure 4. Visually, Mg deficiency in vines is seen as
chlorosis of leaves, especially older ones and cause premature abscission (Abo El-Ezz et al. 2022).
Also, increase the risk of tendril atrophy. Collectively, chlorosis is caused either by Mg, N, P, Mn
Figure 3. Leaves with potassium deficiency symptoms with the reddening progressing from the leaf margins.
JOURNAL OF PLANT NUTRITION 11
and/or Fe deficiency, high content of soil calcium (calcareous soils) or a combination of these fac-
tors. Foliar Mg supplied by the magnesium nanoparticles formulation (Mg-NPs) at 0.5 g/l during
four stage growth (flowering, fruit set, v
eraison and harvesting) compensated Mg deficiency in
the vine, increased chlorophyll content, increased the vine leaf area, improved yield, and the
bunch quality (Abo El-Ezz et al. 2022). Magnesium deficiency in the vineyard, also occurs in soil
with low pH and phosphorus values.
Influence of calcium on the vineyard
Calcium (Ca) maintains fruit firmness as interacts with the cell walls and binds pectin and forms
bridges between pectic acid. It provides firmness, thereby increasing grapesshelf life and protect-
ing them against disease and insect susceptibility. Calcium may play a role in the structure of the
vine and may be associated with bunch necrosis. Calcium deficiency has been reported to cause
incomplete flowering, densely branched shortened roots, chlorosis, and necrosis (Duan et al.
2022). Calcium may be applied to the soil as lime or gypsum. Foliar calcium application (water-
soluble calcium fertilizer (Ca(NO
3
)
2
4H
2
O)) increases sugar, anthocyanins, and total phenol.
Application of water-soluble Ca by drip irrigation increased total phenol, lowered TA, signifi-
cantly decreased the sugar-to-acid ratio and reduced TSS by 6-14% compared to control (Wang
et al. 2019). Similarly, foliar Ca at 2% (w/v) as CaCl
2
for three applications throughout the fruit-
ing season; first spray at the pea size berry and third application a week before harvest results in
low anthocyanins and increased grape berry flavonols, flavan-3-ols, and stilbenoid (Martins et al.
2020). Whereas, the main metabolites E-resveratrol, E-piceid, and E-viniferin increased signifi-
cantly in grape berries with foliar calcium application for enhanced grape quality.
To obtain optimal effect on yield and grape quality, Wang et al. (2019), recommended 30 kg
Ca/ha application by drip-irrigation. Moreover, it was established that bio-regulators (foliar appli-
cation) increased the harvest of grapes, especially in the first and third years of treatment depend-
ing on the preparation and grape cultivar, by 27-64% and 24-29%, respectively as compared to
control. Similarly, fertigation at a rate of 75 kg Ca/ha, increased the yield by almost 30.92% of
wine grapes in Ningxia, China (Wang et al. 2019). Combined foliar application of 1% CaSO
4
and
Table 2. Types of Potassium fertilizers and their corresponding potassium oxide percent.
K fertilizer type Chemical formula %K
2
O
Potassium sulfate K
2
SO
4
48-53
Potassium chloride KCl 60-63
Potassium nitrate KNO
3
 44
Potassium carbonate KCO
3
 25
Potassium magnesium sulfate K
2
SO
4
.2MgSO
4
22
Potassium thiosulfate K
2
S
2
O
3
21
Need to determine nitrogen requirement before application as also supplies N.
For best results used on acidic soil.
For best results used on alkaline soil.
Figure 4. Leaves with magnesium deficiency symptoms.
12 A. JAMES ET AL.
0.5% ZnSO
4
during spring before the bud wooly stage, significantly improved frost tolerance
through a delayed budburst for nine days (Karimi 2019). Delayed budburst was the consequence
of accumulated abscisic acid (ABA), a dormancy hormone in vine treated with Zn and Ca.
Sulfur (S)
Sulfur is an important nutrient required for grapevine physiological growth and development. It
helps to form proteins, coenzymes and chlorophyll, plays a role in energy metabolism, and pro-
mote root growth and seed formation (Considine and Foyer 2015). It is used in the formation of
iron-sulfur complexes of the electron transfer chain during photosynthesis. Symptoms of S defi-
ciency are similar to nitrogen deficiency, yet are rare given the widely adopted use of sulfur-based
sprays for management of downy and powdery mildew and sulfur-containing fertilizers in most
vineyards globally (Ferreira et al. 2022; Savocchia et al. 2011).
Micronutrients
Micronutrients are essentially important as macronutrients to have better plant growth, yield and
quality. Various physical and metabolic functions are governed by these mineral nutrients in the
grapevine. They are required only in trace amounts, which are partly met from the soil or
through chemical fertilizers or other sources. For instance, Zn and Cu may accumulate in the
vineyard soil and grapevine plant structure as the result of consistent use of copper and/or zinc-
based fungicides (Hummes et al. 2019; Korchagin et al. 2020). The most important factor control-
ling soil micronutrients availability for plant utilization is soil pH. In addition to the pH reaction
of the soil to form an aqueous, the presence of carbonates (CaCO
3
) in the soil, low soil aeration,
soil compaction, low organic matter content, low temperature in the root zone, poor biological
properties, and inadequate soil N status are also important factors controlling the supply of
micronutrients for plant nutrition (Likar et al. 2015; Romic et al. 2012; Serrano et al. 2017).
Moreover, micronutrient deficiencies are exacerbated by intensified agricultural practices, unbal-
anced fertilizer application, and depletion of soil nutrients.
Zinc (Zn)
Zinc is an essential nutrient for normal plant growth and development, as is a structural element
of proteins and enzymes in living organisms. Zinc involves in protein synthesis, some hormone
production and fruit set. The work of I~
nigo et al. (2020), presented the values of zinc ranging
from 5 to 136 mg/kg in most vineyards globally. A high concentration of Zn in the soil indicates
correct management practices such as the application of organic manure or sewage sludge.
Moreover, Zn has been studied to reduce the effect of environmental stress by maintaining the
integrity of cellular membranes and decreasing ionic leakage in Zn treated plants. Zinc in the soil
may be accumulated from the consistent use of fungicides containing Zn, such as Mancozeb
(Brunetto et al. 2014).
Zinc deficiency may result in stunted growth and development of small, undersized leaves
with mottling between veins, clawed margins, and widened petiolar sinus. Poor fruit set and hen
and chickenbunches of variable sized berries may occur even when leaf symptoms are not
observed. As the result, pre-flowering Zn foliar sprays are common practice in most vineyards
(Ali et al. 2021). In a combined treatment of 1% CaSO
4
and 0.5% ZnSO
4
, Zn was shown to be
effective in preventing photosynthetic pigment when vines were subjected to stress conditions
(Karimi 2019).
JOURNAL OF PLANT NUTRITION 13
Iron (Fe)
Iron is a micronutrient present in proteins for energy transfer in assimilation and respiration and
involves in chlorophyll formation. Therefore, Fe is important for chlorophyll production, photo-
synthesis, enzyme composition, energy transfer as well as N fixation and reduction. In the soil, Fe
originates from complexes of soil inorganic minerals and organic matters. Iron is largely present
in an inorganic form, predominantly as amorphous Fe, goethite, hematite, and ferrihydrite.
Annual removal of Fe in the vineyards is relatively low and the total amount in the soil would
not justify the development of Fe deficiency, which often occurs as a result of poor availability of
Fe for the vine. Iron availability is usually restricted by bicarbonate in the calcareous, poor soil
aeration, and compaction or waterlogged alkaline soils and low level of soil organic matter
(Tagliavini and Rombol
a2001). Another peculiar aspect of Fe deficiency in the vineyard is related
to the size and age of the vines and to the fact that after absorption Fe has to be transported for
a long distance to reach the canopy.
The deficiency of Fe causes stunted growth, diffuse yellowing of shoot tips and young leaves as
well as reduced fruit yield and quality. Severe Fe deficiencies cause the whole leaf to become
chlorotic, while the vein remains green or sometimes atypical uniform chlorosis and necrosis.
Consequently, when chlorotic symptoms in the vineyards develop, fruit yield and quality can be
severely reduced in the current season as well as next season fruiting as fruit buds poorly develop.
Foliar Fe application has been reported to effectively treat Fe deficient syndromes and chlorosis
in grapevines (Zebec et al. 2021). Other management such as the use of organic manure to
enhance soil organic matter and improve soil aeration, which may prevent the re-crystallization
of ferrihydrite.
Copper (Cu)
Copper is the component of enzymes involved in oxidation as well as chlorophyll synthesis.
Deficiency symptoms are not common in most vineyards, although may be expressed as low
vigor, shoots do not mature, barks appear rough, hence poor production. Copper in soil may be
accumulated from consistent use of copper-based fungicides such as Bordeaux mixture (Ca(OH)
2
þCuSO
4
) and Cu-oxychloride (CuCl
2
.3Cu(OH)
2
) (Brunetto et al. 2014). Copper toxicity results
in decreased levels of other essential elements like P, Fe, and Zn, which has been reported in
areas of persistent copper fungicide use. The values of Cu in the vineyard soil were presented by
I~
nigo et al. (2020), which ranged from 3.01 to 178.1 mg/kg.
Manganese (Mn)
Manganese plays a major role in nitrogen metabolism and the synthesis of chlorophyll.
Manganese is present in the soil as exchangeable manganese or manganese oxide. Mn uptake
increases the photosynthesis, assimilation and relative growth and yield. Also, it is involved in
defence mechanisms, acting as a co-factor in many enzymes involved in the biosynthesis of lignin
and flavonoids as well as superoxide dismutase (SOD), which defends the plant cells against free
radicals. Mn reduces water stress and helps vines to grow under water, salt or high, and low-tem-
perature stresses. Foliar application of 2 or 4 g Mn/l (MnSO
4
) on the grapevine ameliorated the
effect of drought as indicated by plant height, root length, leaf surface area, total chlorophyll con-
tent, and number of leaves per plant (Ghorbani et al. 2019). Moreover, carbohydrate, proline, and
protein contents were higher in treatment than in the control. Manganese deficiency is expressed
as yellowing of the interveinal area of older leaves and may be mistaken for zinc or Fe deficiency.
The symptoms are pronounced in vines on sandy, calcareous soils or in areas of high rainfall.
Toxicity of Mn is rare in vineyards but can be seen as black spots on the leaves and bunch stems.
14 A. JAMES ET AL.
Boron (B)
Boron exists in the soil as anion borate, though availability for the vine is limited due to high
solubility and leaching by rain or irrigated water in shallow or coarsen textured soil. It plays a
role in the synthesis of hormones regulating plant growth, pollen tube growth, flowering, and
fruit set, carbohydrate translocation such as sugar, as well as active salt absorption (Parthiban
et al. 2021; Tadayon and Moafpourian 2019). Also, B is essential for the development of plant
meristem and metabolism of amino acids, proteins, carbohydrates, Ca, and water. It contributes
to better utilization of Ca in plants S et al. (2021). Thus, when the plant is B deficient, it cannot
utilize Ca properly. Whereas, B accumulation in plant increase with increasing soil K. Boron ferti-
gation improves the synthesis and movement of carbohydrates, especially sucrose from the leaves
to the roots and fruiting bodies (Bredun et al. 2021). However, excessive B may lead to the accu-
mulation of sugar in the leaves.
Boron deficiency shows various symptoms including stunted growth, such that vines interno-
des are shortened displaying a zig-zag pattern, meristem distortion, death of shoot tips, intervei-
nal chlorosis of older leaves and poor flowering (Peacock and Christensen 2005). In case of
several B deficiencies, bunch and tendril abortion can occur affecting pollen tube growth, hence
poor fruit set, small short berries, and fruit cluster necrosis. Boron toxicity symptoms include
poor roots development, cupped leaves on young shoots, followed by brown necrotic spots on the
leaf margin and yellow streaks between veins, which then merged and result in decreased vine
vitality (Christensen, Beede, and Peacock 2006). For a better outcome, a direct spray of B on the
vine should be before the flowering stage to achieve productivity, maturation, and chemical com-
position of grapes (Christensen, Beede, and Peacock 2006; Tadayon and Moafpourian 2019). For
instance, foliar spray of B as boric acid (1, 2 or 4 g/l) during five phenological phases; end of
bloom, buckshot berries, pea-size berries, v
eraison, and 15 days before harvest has resulted in
increased yield and productivity, berry TSS, pH, and improved berry texture as well as cell wall
integrity (Bredun et al. 2021).
Silicon fertilizers (Si)
Silicon (Si) is the second most abundant element in the earths crust and soils. However, the
plant-available form of Si is usually present in low amounts because of the extremely low solubil-
ity of aluminosilicate clay minerals. Therefore, the application of Si fertilizer has been a rather
common agricultural practice worldwide. Silicon has been studied to strengthen cell walls,
improve plant strength and health and boost the vine to fight pests and diseases. It improves root
as well as plant biomass, crop yield, and productivity (Bassiony and Ibrahim 2016). Several stud-
ies have shown that silicate fertilizers not only significantly provide improved development,
growth, and increased crop yield, but also, largely improve crop quality parameters such as fruit
sugar, vitamin C, and phenolics. Also, Si induces plant tolerance to a wide range of abiotic and
biotic stress including tolerance to drought or frost and resistance against pathogens, which may
hamper grape growth (Gomes et al. 2020). For instance, Si fertilizers increased grapes yield by
13.5%, improved TSS, the ratio of TSS/TA and fruit firmness at the rate of 600 kg Si fertilizer
(SiO
2
)/ha (Zhang, Liang, and Chu 2017). Moreover, the application of 4 and 8 g Si/l as sodium
metasilicate and 15 g Ca/l as calcium chloride per grape plant represent a potential alternative fer-
tilizing strategy, which could increase yield without damaging the phytochemical characteristics of
the fruit and its derivatives (Gomes et al. 2020). While, the treatment effect of 4 g Si/l and 5 or
15 g Ca/l led to a high yield per plant and high productivity compared to control (Gomes et al.
2020). Moreover, Bassiony and Ibrahim (2016), applied 0.1% Si (as 1 g SiO
2
in 100 ml of KOH
0.1 N solution) combined with 2% Moringa leaves extract every two weeks after 15-20 cm shoot
length to v
eraison resulting in enhanced vegetative growth, leaf nutrients content as N, P, K, Fe,
JOURNAL OF PLANT NUTRITION 15
Zn, and Mn, and must quality in terms of SST, anthocyanins, and total sugar in the
Egyptian vineyards.
Molybdenum (Mo)
Molybdenum is part of the nitrate reductase and nitrogenase enzymes, which involves in nitrogen
metabolism and are used by the plant to reduce nitrates into usable forms (Kaiser et al. 2005).
Mo is found in soil as molybdate (MoO
42-
) and its availability is pH dependent, enhanced in
alkaline soils. Molybdenum is present in as many as different complexes depending on the chem-
ical specification of the soil. Mineral forms of Mo include molybdenite (MoS
2
), ferrimolybdenite
Fe
2
(MoO
4
)
3
, and wulfenite (PbMoO
4
) (Sun and Selim 2020). The vine Mo deficiency symptoms
include stunted growth, abnormal flower formation, and abortion associated with low yield and
increased susceptibility to low temperature and waterlogging stress (Longbottom, Dry, and
Sedgley 2010; Rana et al. 2020). Also, it has been suggested that low-temperature stress in vine
may be due to the breakdown of Mo-containing enzymes. Particularly young merlot vines show
vegetative symptoms such as zig-zag shoot growth, inky epidermis, and rolled leaves with burn
margins. In Merlot causes poor fruit set as Merlot problem, however foliar Mo applications have
been used to successfully improve Merlot growth and productivity (Longbottom, Dry, and
Sedgley 2010). Molybdenum in various forms, particular foliar sodium molybdate has been
applied to grapevines as a remedial measure to supplement Mo deficiency, rescue the activity of
molybdoenzymes, and combat low yield (Kaiser et al. 2005).
Conclusion
In this review, we found different fertilizer ratios of N, P, K, and micronutrients foliar application
have a significant impact on yield and grape berry quality. Balanced application of N increases
yield and YAN in must. Equally important, application of organic manure gradually release
nutrients for the vine, increase soil water holding capacity, improve soil texture and organic mat-
ter content, and create a nutrient-rich ecosystem. The overall result of fertilizer application in the
vineyard increases the number of bunches per vine, bunch length, berry number per bunch, berry
diameter, and bunch weight, leading to increased yield and improved grape quality. Thus, to
establish an effective fertilization program, the decision-making process should have information
on target yield and vineyard management strategies influencing the efficiency of the use of
nutrients present in leaves and canes. Soil testing and plant analysis continue to be of paramount
importance; and integrating all possible data will help to decide on fertilizer rates to ensure the
adequate nutritional status of the grapevines, without compromising the quality of wine and the
environment. Therefore, rational use of fertilizers and nutrient-specific fertilization plans are rec-
ommended for the vineyard.
Acknowledgement
We would like to thank everyone who has supported us to reach our goals.
Disclosure statement
No potential conflict of interest was reported by the authors.
ORCID
Athuman Mahinda http://orcid.org/0000-0001-6752-6195
16 A. JAMES ET AL.
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22 A. JAMES ET AL.
... Both systems have their advantages and disadvantages. Organic fertilization is environmentally friendly, but it is slow-acting and may contain low concentrations of some essential nutrients, whilst inorganic fertilization, on the other hand is fast-acting, provides nutrients in readily available forms, and cost-effective (James et al., 2022). ...
... Additionally, fertigation systems have been shown to significantly influence plant growth, yield, photosynthesis, and secondary metabolites (Incrocci et al., 2017;James et al., 2022). Properly timed and applied fertigation can provide plants with the necessary nutrients they need for optimal growth and development, leading to increased yields. ...
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Nowadays a balance in plant production is required, especially in terms of nutrition, yield and for an optimal aroma and sensory profile of resulted wines. In this research, we compared two different methods of plant nutrition: one-single application (single-fertilization-SF) and two applications with the same amount adopting the fertigation (split fertigation-SpF), including a control not-treated, in a specific Italian region where grapevine Schiopettino cv. is being cultivated and this practice was not investigated yet. SpF promoted the photosynthesis parameters compared to a SF and to the non-treated vines (NTC). On these basis, the biological and physiological activity of the whole plant was enhanced. SpF treatment tendentially and significantly improved the qualitative, productive, physiological, and oenological parameters of "Schioppettino" wine.
... It creates favorable environments for fungal diseases such as downy and powdery mildew, and bunch stem necrosis which ultimately reduce grape yield [46]. The reduced light interception in the fruit zone as a result of higher canopy density delays fruit ripening and lowers the content of total soluble solids (TSS), anthocyanins, terpenes, and total phenols in grapes [47]. The application of N on grapevines should be undertaken with caution because of its contradicting effects on vegetative growth, yield, and chemical composition of the grapes, must and wine. ...
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Grape (Vitis Vinifera) is among the horticultural crops which are consumed as fresh or processed products due to its nutritious importance. High yield and quality grape production are the major focus of grape producers and stakeholders in the value chain. However, nutrient availability is a challenge to reach the targets which cause deficiency of some essential elements for plant uptake. Fertilizer application, either organic or inorganic supports grapevine development accounting for the final yield and quality grape. Organic fertilizer for instance, farmyard, compost and/or vermicompost manure in grapes improves physical and biological properties of the soil by decomposition which makes major nutrients readily available for plant absorption. Supplementation of inorganic fertilizer in grapevines increases macronutrients and micronutrients which enhance plant growth and reduce nutrient deficiencies. Moreover, it provides essential elements required for wine making to produce quality wine. Nevertheless, the application of either organic fertilizer or in organic fertilizer alone has either positive or negative effects in grape production. Therefore, this review discusses the role of organic fertilizer, inorganic fertilizer, essential nutrient elements and the importance of integrated use of organic and inorganic fertilizer as the effective approaches for the production of grape and quality wine.
... Excessive use of chemical fertilizer can lead to the accumulation of nitrates in vegetable products-this negative impact can be mitigated by reducing the amount of chemical fertilizer used and combining them with organic fertilizer that have better nutritional properties [18][19][20]. In grape production, the application of organic fertilizers ensures the supply of organic carbon, improves the organic matter content in the soil and promotes soil microorganisms [9,21]. The combined use of compost, mineral fertilizers and microbial agents promotes root development and improves soil quality in table grapes [22]. ...
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Increasing the use of organic fertilizers is an effective measure to improve, increase soil fertility and maintain crop yields. The aim of this study was to investigate the influence of different types of organic fertilizers on the early yield of grapes and soil parameters, as well as the relationship between soil parameters and grape yield under fertilization conditions. The ‘Shine Muscat’ grape was used as the material, with early maturing cultivation in the solar greenhouse. From the time of grape planting, three-year continuous fertilization management was carried out using five types of base fertilizers: chemical fertilizer (CK), fermented corn stalk residue (A1), mature sheep manure (A2) and two types of commercial organic fertilizers (B1 and B2). In the third year, berry and soil samples were collected to determine grape yield and evaluate soil physicochemical properties, nutrient status and changes in enzyme activity, studying the relationship between grape yield and soil indicators. The results show that compared to CK, the grape yields with B1 and B2 increased by 19.04% and 16.26%, respectively, while A1 and A2 decreased by 24.09% and 18.97%. Organic fertilizer application reduced soil bulk density, increased soil porosity, enhanced soil organic matter content and effectively buffered soil pH levels. Two types of commercial fertilizers (B1 and B2) improved soil total nitrogen, total phosphorus, total potassium, available nitrogen and available phosphorus content to varying degrees. All organic fertilizer treatments effectively increased soil enzyme activity, except for soil sucrase activity. Through correlation and regression analyses, it was found that in this study, the levels of available nitrogen, available potassium and soil saccharase activity were most closely related to early grape yield. Their influence on grape yield was in the order of available nitrogen > soil saccharase activity > available potassium. Therefore, with a combined organic and inorganic basal fertilization system, the fertilizer nutrients are more comprehensive and help to increase the productivity of grapes at the beginning of the fruiting period. Nitrogen and phosphorus are very important in the maintaining of grape yields and improving sucrose activity in the soil through the application of organic fertilizers cannot be ignored when increasing yields.
... In the context of vineyard production, AMF species play a crucial role in water uptake, aiding the host plant in resisting abiotic and biotic stresses without compromising essential characteristics for vine production (Nerva et al., 2022). When we initiated this study in a vineyard field in Southern Brazil, we did not anticipate discovering strong AMF-host pairing specificity Liu et al., 2022;James et al., 2022). Thus, AMF species are emerging as a key factor in this context. ...
... Mineral nutrition plays a pivotal role in grape production, exerting influence over all aspects of vine growth, berry development and maturation, and ultimately, wine quality [310,311]. The primary macronutrients, i.e., N, P, and K, control critical grapevine metabolic processes, which influence grape quality and thus the eventual alcoholic fermentation process [46]. ...
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Soil health encompasses the effects the uppermost part of the land have on human wellbeing in a broad sense, because soil is where most food ultimately comes from, and because it more inconspicuously fulfils other ecological functions, as important as feeding, for our planet’s welfare, which is ours. Viticulture exploits the soil’s resources from which wine, its most valuable produce, boasts to obtain some of its unique quality traits, which are wrapped within the terroir concept. However, using conventional methods, viticulture also has harsh impacts on the soil, thus jeopardizing its sustainability. How long will the terroir expression remain unchanged as vineyard soil degradation goes on? While this question is difficult to answer because of the complex nature of terroirs, it is undeniable that conventional soil management practices in viticulture leave, in general, ample room for improvement, in their impact on vineyards as much as on the environment. In response, viticulture must adopt practices that enable the long-lasting preservation of its grounds for both on-farm and off-farm benefits. In this regard, the increase in the soil’s organic matter alongside the enhancement of the soil’s biological community are key because they benefit many other soil properties of a physical, chemical, and biological nature, thus determining the soil’s healthy functioning, where the vines may thrive for a long time, whereas its surroundings remain minimally disturbed. In the present review, the importance of soil health as it relates to vineyards is discussed, the soil degradation factors and processes that threaten winegrowing areas are presented, successful soil-health enhancement practices are shown, and future research trends are identified for the benefit of researchers and stakeholders in this special agricultural industry.
... For instance, chlorosis is often a physiological symptom, not a disease in itself. It is caused by factors such as nutrient deficiencies (iron, nitrogen, magnesium, or zinc), poor soil conditions (inadequate drainage, high pH, or compacted soil), or environmental stress (water imbalance, root damage, or exposure to pollutants), all of which lead to the yellowing of leaves [5,6]. The most prevalent method of mitigating biotic stress in grapevines involves the administration of chemical substances. ...
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Changes in the soil fertility, physiological traits, and fruit quality of Morinda citrifolia were investigated between January 2012 and December 2020. Soil samples, physiological properties, and fruits of M. citrifolia were collected in a factorial scheme 2 × 2 × 2, with the use of intercropping with Musa spp., mulching, and biofertilization (e.g., presence and absence) within five blocks. We evaluated soil reaction (pH), P by Melich-1, soil exchangeable cations, potential acidity (H+ + Al3+), sum of bases, CEC, SOC, Fm, F0, Fv, Fv/F0, Fv/Fm, photochemical quenching, non-photochemical quenching, apparent electron transport rate, firmness, refractometric index, soluble sugar content, acidity, ascorbic acid, total phenolic content, antioxidant activity, and fruit mineral content. Our results emphasized the influence of agroecological practices on soil chemical properties, plant physiological traits, and fruit quality of M. citrifolia. The combined use of intercropping, mulching, and biofertilization showed high values of soil pH, available P, Mg2+, electron transport rate, firmness, ascorbic acid, antioxidant activity, fruit Zn content, and fruit Na content and low values of Ca2+ and K+ under field conditions. The results of this study highlight the importance of considering agroecological practices as promoters of fruit quality, and thus improving fruit properties with high importance for human mental health, fruit palatability, fruit development, fruit postharvest storage, and redox state of the human body.
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Bağcılıkta en önemli hedeflerden biri omcalardan yüksek verim elde etmek ve dolayısıyla birim alanda üretilen yaş üzüm miktarını arttırmaktır. Bu amaca yönelik bağlarda pek çok kültürel uygulama yapılmakta ise de omcalara uygulanacak optimum budama düzeyinin belirlenmesi özel bir öneme sahiptir. Çünkü asmaların bir yaşlı dalları üzerinde yer alan kışlık gözleri, salkımları oluşturacak olan yazlık sürgünlerini taslak halinde taşımakta ve üzüm çeşitlerinin yüksek verimliliğe sahip göz düzeyleri birbirlerinden farklılık gösterebilmektedir. Bu çalışma, Vitis vinifera L. türüne ait on iki üzüm çeşidinin göz verimliliklerinin salkım teşekkül pozisyonları dikkate alınarak belirlenmesi amacıyla 2022-2023 yıllarında yürütülmüştür. İncelenen tüm üzüm çeşitlerinde, sürgünler üzerinde yer alan ilk 10 gözün verimlilikleri birbirlerinden farklı bulunmuştur. Bununla birlikte, belirli bir göz pozisyonunun verimliliği bakımından üzüm çeşitlerinin birbirlerinden farklılık gösterdiği belirlenmiştir. Dahası, salkım teşekkül pozisyonları incelenerek bir üzüm çeşidinin göz verimliliğinin belirlenebileceği saptanmış ve bu yöntemin pratikte uygulama kolaylığı sayesinde üreticilere tavsiye edilebileceği kanaatine varılmıştır. Elde edilen bulgular ışığında; Şiraz üzüm çeşidinin 4. göz üzerinden, Trakya İlkeren, Yalova İncisi, Alphonse Lavallée ve Boğazkere üzüm çeşitlerinin 4. veya 5. göz üzerinden, Perlette, Red Globe, Banazı Karası ve Öküzgözü üzüm çeşitlerinin 5. göz üzerinden, Flame Seedless, Royal ve Italia üzüm çeşitlerinin ise 5. veya 6. göz üzerinden budanması gerektiği sonucuna varılmıştır.
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Green pruning practices are frequently used in viticulture. These practices provide better light penetration, successful air circulation, prevents air humidity and aids disease control in the grapevine canopy. However, excessive leaf removal inhibits vine growth and good fruit ripening. In this study, the effects of some foliar and canopy management practices on cluster, vegetative growth traits and bud fruitfulness of the following year were investigated. Some green pruning (unproductive shoot removal, leaf removal, shoot tip removal, shoot topping), foliar microelements and green pruning + microlements treatments were applied in ‘Trakya İlkeren’ grape cultivar. Microelement application increased cluster (382.1 g) and berry weight (4.4 g), berry length (19.7 mm) and width (18.2 mm), cluster length (19.5 cm) and width (13.9 cm), berry volume (3.30 cm3) and berry firmness (7.46 N). The highest maturity index was detected in the control group (23.4). While the most intense L* color value was determined in the green pruning + micro element application, the b* color value was determined in the green pruning application. It was determined that leaf area (211.2 cm2), leaf chlorophyll index (32.75 SPAD), shoot diameter (11.13 mm), shoot length (194.71 cm) and internode length (76.81 mm) increased with microelement application. Among the treatments, the most effective application on bud fruitfulness in the following year was at microelement application (1.34 clusters). The effects of the treatments on the amount of Total soluble solids pH, titratable acid and a* color value were not significant. In this study; green pruning + microelement application was recommended for yield and quality sustainability. According to bud fruitfulness results, the highest cluster numbers per node were obtained from pruning with 4 to 10 buds. It is recommended to prune medium or long to obtain higher yields from this cultivar.
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Nitrogen nutrition is one of the most effective cultural practices in vineyards. The vine nitrogen status influences the berries' quality characteristics and the produced wines. The current study investigated the effect of traditional nitrogen fertilization in the form of ammonium sulfate compared to nitrogen fertilization coupled with the nitrification inhibitor 3,4-Dimethylpyrazole phosphate (DMPP) on the agronomic characteristics of grapes and the produced wines of the white variety Savvatiano from a productive vineyard in the Attiki region. Must and wine quality was evaluated by a chemical analysis and sensorial evaluation by trained panelists. The different forms of nitrogen fertilizers did not significantly affect the aroma and sensory profile in contrast to unfertilized grapevines. In addition, the applied fertilization increased some important aroma compounds in the wine, compared to no fertilization. The significance of this work is to add information about the effect of nitrogen fertilization on the wine volatile composition of the Greek white grapevine Savvatiano.
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The nitrogen (N) critical level in leaves and maximum technical efficiency (MTE) doses contribute to the grape yield, must quality, and fertilizers rationalization. This study aimed to define sufficient ranges and critical levels in leaves and maximum technical efficiency doses to obtain high yields and quality must in grapevines grown in sandy soil in a subtropical climate. ‘Chardonnay’ and ‘Pinot Noir’ grapevines were subjected to the annual application of 0, 10, 20, 40, 60, and 80 kg N ha−1 in a vineyard. The nitrogen concentration in leaves at flowering and veraison, grape yield and grape must chemical parameters (total soluble solids—TSS, total titratable acidity—TTA and total anthocyanins—TA) were evaluated. The N critical level in leaves at flowering was different between grapevines cultivars, but this was not observed in leaves at veraison. It was possible to estimate MTE for ‘Chardonnay’ grapevines, in the evaluated growing seasons. In the range of higher probability of the N critical level in leaves, the TSS and TTA variables showed opposite responses, for both cultivars. This study proposes different N critical levels for red and white grapevines, in a subtropical climate.
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This study investigated the interactions between different irrigation and nitrogen (N) fertilization rates and their effects on the 15N fertilizer absorption and utilization of Cabernet Sauvignon grapevines in a vineyard near Xinjiang, China. The fertilizer treatments consisted of three 15N-labeled urea nitrate fertilizer applications (191.4, 254.4, and 317.4 kg/ha). The irrigation treatments included two fractions (0.75 and 1.0) of estimated vineyard water use (ETc). The results showed that: (i) the residual amount of 15N fertilizer in the soil was mainly in the surface layer (0–20 cm), the residual fertilizer N in the surface layer accounted for 68–87% of the total residual N, the residual 15N fertilizer at different depths differed significantly, and the residual 15N was positively correlated with the amount of fertilizer applied (p < 0.05); (ii) The absorption of fertilizer N by grapevines accounted for only 12–17% of the total N absorption, and the proportion absorbed from soil N was as high as 82–87%. There was no significant difference in the amount of N absorbed between different water and fertilizer treatments; (iii) The 15N uptake under the different water and fertilizer treatments differed and was significantly higher in the roots than in other organs, followed by the fruit, the leaves, and finally, the stems. Our results provide a reference for improving the soil environment and encouraging a sustainable development of the grape industry. According to the experimental results, it was recommended that farmers adopt irrigation levels of 1.0 ETc and fertilizer application of 254.4 kg N/ha.
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Powdery mildew caused by the fungus Erysiphe necator is a major grape disease worldwide. It attacks foliage and berries and reduces yield and wine quality. Fungicides are mainly used for combating the disease. Fungicide resistance and the global requisite to reduce pesticide deployment encourage the use of environment-friendly alternatives for disease management. Our field experiments showed that the foliar application of the potassium phosphate fertilizer Top-KP+ (1-50-33 NPK) reduced disease incidence on leaves and clusters by 15–65% and severity by 75–90%, compared to untreated vines. Top-KP+ mixed with Nanovatz (containing the micronutrients boron (B) and zinc (Zn)) or with TruPhos Platinum (a mixture containing N, P2O5, K2O, Zn, B, Mg, Fe, Mn, Cu, Mo, and CO) further reduced disease incidence by 30–90% and disease severity by 85–95%. These fertilizers were as effective as the fungicide tebuconazole. Tank mixtures of fertilizers and tebuconazole further enhanced control efficacy in the vineyards. The modes of action of fertilizers in disease control were elucidated via tests with grape seedlings, microscopy, and berry metabolomics. Fertilizers applied preventively to the foliage of grape seedlings inhibited powdery mildew development. Application onto existing mildew colonies plasmolyzed mycelia and conidia and arrested the development of the disease. Berries treated with fertilizers or with a fungicide showed a significant increase in anti-fungal and antioxidant metabolites. Twenty-two metabolites, including non-protein amino acids and carbohydrates, known for their anti-fungal and bioactive effects, were significantly upregulated in grapes treated with fertilizers as compared to grapes treated with a fungicide, suggesting possible indirect activity against the pathogen. Esters and organic acids that contribute to wine quality were also upregulated. We conclude that integrating macro and micronutrients in spray programs in commercial vineyards shall control powdery mildew, reduce fungicide deployment, delay the buildup of fungicide resistance, and may improve wine quality.
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This study characterized growth characteristics and cellular details employing microscopy techniques in hydroponically-grown Ca2+-sufficient and Ca2+-deficient grapevines (Vitis vinifera) in a glasshouse. The Ca2+-deficient vines exhibited significant reductions in shoot length, shoot and trunk fresh weights, leaf area, chlorophyll, which eventually led to drooping, yellowing, and chlorosis of leaves. Roots were less dense and primarily dark and necrotic. Furthermore, their xylem vessels were small, polygonal, and appeared to be collapsed yet increased in number and developed lateral roots. Despite such alterations, the anatomical organization of leaves was not affected, yet they developed with more xylem vessels with thick walls and lignin in their mesophyll and vascular tissues. The chloroplasts in internodes’ chlorenchyma, phloem, and cambium underwent significant ultrastructural modifications. The concentrations of macro and micronutrients varied significantly among the roots, trunk, canes, and leaves, including the growth characteristics. These structural and growth modifications of calcium deficiency enable us to understand better the link between the symptoms and functions and for a holistic understanding of Ca2+ functionalities.
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