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Mendel University in Brno, Czech Republic Faculty of Horticulture
Mendel University in Brno, Czech Republic Faculty of Horticulture
FACULTY OF HORTICULTURE
DISSERATION
In partial fulfillment of the requirements of the degree of
Doctor (Ph.D.)
By
Ing. Mohamed Ewis Abdelaziz
Supervisor
Doc. Ing. Robert Pokluda, Ph.D.
ABSTRACT
In two growing seasons of 2006 and 2007, an experiment was carried out to study the
effect of inoculation with Azotobacter chroococcum, Azospirillum brasilense, Glomus
mosseae and their combinations on growth, yield and selected component composition of two
cucumber cultivars (Cucumis sativus L. cultivars ‘Passandra’ and ‘Girola’) grown on
rockwool and peatmoss under an open soilless system in greenhouse at Faculty of
Horticulture, Mendel University, Czech Republic. Vegetative growth characteristics, number
of fruits per plant and fruits dry matter of cultivar Passandra were significantly higher than
cultivar Girola. On contrast, early and total yield did not reveal much variation between
cultivars. Nitrogen in fruits of cultivar Passandra and P, K, Ca in fruits of cultivar Girola
revealed higher significant increases in the first season while N, P, and Mg did not differ in
leaves of the two cultivars, with some exceptions. However, type of cultivar or substrate did
not affect vitamin C or TSS contents. Nitrate, recorded higher significant increase in fruits of
cultivar Girola, but values still under the critical limits. Plants grown on peatmoss revealed
significant increase in number of leaves/plant and leaf DW. Moreover, light significant
increase was obtained in early yield, fruit length and weight of cucumbers grown on
st nd
rockwool. All inoculation treatment improved total yield by 8 and 21 % in 1 and 2 seasons,
respectively. In addition, significant increase in average fruit weight was found when plants
st
inoculated with Azotobacter alone or combined with Azospirillum than control in the 1
season. Microorganisms inoculation improved the marketable yield, mineral component in
leaves and fruits of cucumbers, with some exceptions. Vitamin C recorded the highest
significant increase with Glomus + Azospirillum. Furthermore, N-Fixers alone/or combined
with Glomus improved growth of cucumber plants grown in peatmoss. In particular, growth
and yield of cultivar Passsandra grown in peatmoss were promising responded to
microorganisms inoculation than other treatments under the open soilless culture.
V letech 2006 a 2007 byly provedeny experimenty studující vliv inokulace Azotobacter
chroococcum, Azospirillum brasilense, Glomus mosseae a jejich kombinací na růst, výnos a
vybrané obsahové složky okurek Cucumis sativus L. (odrůdy ‘Passandra’ a ‘Girola’)
pěstovaných v čedičové plsti a v rašelině v otevřeném hydroponickém systému ve skleníku
Zahradnické fakulty v Lednici, Mendelovy Univerzityv Brně, Česká Republika.
Vegetativní růstové charakteristiky, počet plodů na rostlinu a obsah sušiny plod ů byl
průkazně vyšší u odr ůdy Passandra. Na druhé straně ranost a celkový výnos nebyl odr ůdou
ovlivněn. Obsah dusíku v plodech odrůdy Passandra a obsah P, K, Ca v plodech odrůdy
Girola byl průkazně vyšší v první sezón ě. Na druhé straně obsah N, P a Mg v listech byl mezi
odrůdami, až na n ěkolik výjimek, srovnatelný. Podobn ě genotyp neovlivnil obsah vitaminu
C ani TSS. Obsah dusičnanů byl vyšší u odr ůdy Girola, ale stále pod hygienickým limitem.
Rostliny pěstované v rašelinovém substrátu vykazovaly vyšší počet listů na rostlinu a vyšší
zastoupení sušiny v listech oproti variant ě v čedičové plsti. Ovšem čedičová plsť přispěla k
průkaznému, byť mírnému, zlepšení ranosti, délky a hmotnosti plodů okurek.
Všechny inokulované varianty vykázaly vyšší celkový výnos o 8 a 21 % v 1. a 2. pokusném
roce. Také byl zjištěn průkazný nárůst hmotnosti plodů při inokulaci rostlin Azotobacter
samotným nebo v kombinaci s Azospirillum, ve srovnání s kontrolou. Inokulace také zvýšila
tržní podíl výnosu, zastoupení minerálních látek vlistech a plodech okurek, i když s n ěkolika
výjimkami. Obsah vitaminu C byl pr ůkazně zvýšen ve variant ě ošet řené Glomus +
Azospirillum. Vedle toho N-fixujíxí bakterie samostatně nebo v kombinaci s houbou Glomus
podpořily růst okurek pěstovaných v rašelinovém substrátu. Obzvláště růst a výnos odr ůdy
Passandra v rašelinovém substrátu ukázal pozitivníodpověď na inokulaci ve srovnání s dalšími
sledovanými variantami v otev řené hydroponické kultuře.
Page
ABSTRACT ii
I. INTRODUCTION 1
II. REWIEW OF LITERATURE 3
II .1. Crop 4
II. 2. Effect of substrate 6
2.1. Type of substrate 9
2.1.1. Rockwool 10
2.1.2. Peatmoss 11
2.2. Vegetative growth characteristics 14
2.3. Yield and its component 15
2.4. Chemical components 17
II .3. Effect of microorganisms 19
3.1. Nitrogen fixing bacteria 19
3.2. Arbuscular mycorrhiza 23
3.3. Vegetative growth characteristics 26
3.4. Yield and its component 29
3.5. Chemical components 30
III. MATERIALS AND METHODS 32
Thus, water uptake, buffering and availability as well as oxygen diffusion must be
optimal. Also pH and concentration of nutrients and other salts should meet the plant demands
as well as biological activity should be such kind does complete with the plants (Verhagen,
2009). Therefore, the bellow factors will be reviewed:
1. Crop
3. Microorganisms inoculation
3. Mineral components
II. 1. Crop
Cucumbers (Cucumis sativus) belong to the same family, Cucurbitaceae, as
watermelon, zucchini, pumpkin, and other types of squash. Greenhouse cucumber production
is very popular in many areas of the world. The cucumber is a warm season crop with
o
required growing conditions of 27 to 30 C and plenty of sunlight.
A fruit may be developed at each node and more than one fruit may begin to develop
at some nodes. It is usually best to thin these multiple-fruit clusters to a single fruit; however,
vigorous plants can sometimes mature more than one cucumber at a node. Miniature and beit-
alpha types will support several fruit per node. Any distorted fruit should be removed
immediately. The greatest growth of the fruit occurs between day 6 and 14 after the bloom
opens (anthesis). Maximum fruit length occurs at day 14 followed by diameter increase.
(Hochmuth, 1990)
Cultivars of cucumber are grown either to be eaten fresh or to be pickled. Those that
are to be eaten fresh are commonly called slicing cucumbers. Cucumbers such as gherkins
that are specially cultivated to make pickles are oftentimes much smaller than slicing
cucumbers.
Accurate statistics on the acreage of greenhouse devoted cucumber production are not
easily obtained as no official accounting is made by any government or private organizations
(Benton, 2005).
Quality of growing media is largely defined as the ability to perform various intrinsic
and extrinsic functions, presented by a suite physical, chemical and biological prosperities,
namely to:
(ii) Regulate and partition water flow and storage in the environment
In contrast, the composition of organic growing media, their C/N ratio and
environmental conditions during storage is integral to the growing media biological qualities.
Bohme et al (2008) mentioned that organic substrates show a higher CO 2 outflow than
mineral medium.
The environmental conditions for the microorganisms inhabiting the growing media
are continuously changing, as the microbial density increases and the substrate concentration
simultaneously decreases.
The most dramatic change, however, occurs with the introduction of the plant or the
transplant stage, as a producer of organic carbon is introduced into the system. Root exudates
release a pool of organic carbon (Whipps, 1990) which in turn affects the composition of the
micro flora and microbial mediated processes.
Fig. 1. Major events during the use of growing media in terms of carbon turn over*
GM production site
Introduction
of the plant Treatment
prior to reuse Introduction
of the plant
The influx of root or growing media originating organic carbon and the added water
volume have sustainable impact in the horticultural system. If irrigation is performed at a high
drainage percentage, microbial growth is limited and microbial cells may even be washed out.
In contrast, low dilution rates occur when the added volume water corresponds to the
plant’s water demand or when the carbon influx is low. The first case will lead to an increase
in cell density where as the latter one may cause cell death due to starvation, as limiting
nutrients may not be available fast enough to permit maintenance of cell metabolism. If the
growing media with a plant is considered from a strictly microbiological point of view,
+
substrate limited growth can be maintained as long as essential nutrient factor, such as H
donors, C, N, S or P sources, are available. As long as the actual growth constant (Fuchs,
2006) is lower than maximum at substrate saturated conditions, the dilution rate may be varied
over a large range without washing out microorganisms.
Different investigations show a stability in microbial density over time (Berkelmann,
1992); however, shifts occur with respect to the microbial community structure (Khalil and
Alsanius, 2001; Bergstrand et al, 2009).
Many years ago, Puustjarvi (1997) described a 20-25% decrease of peat volume in
carnation culture. The rapid and extreme variations in water and air availability to roots lead
to frequent cycles of watering (fertigation) and drying (evaporation, root uptake) during
growing management called shrink/swell phenomena (Heiskanen, 1995 and Michel et al,
2004).
-1
Table 2. Bulk density of the substrates used in cucumber production (g.L )*
Stages
Media type New After After
+
first use second use
Perlite 132 151 106
Rockwool 50 70 106
* +
Bohme, et al (2008) with dried roots
This phenomena lead to a change in solid phase and pore organization, which could
then influence the water and air circulation in growing media (Table 2), leading to a decrease
in aeration that could cut off the supply of oxygen to the root during growth (Lemaire et al,
2003).
High irrigation frequency may improve crop performance due to a higher availability
of nutrients, especially P and Mn, while maintaining constantly higher moisture levels in the
root zone.
The only precaution regarding the application of a frequent irrigation schedule is the
possible creation of excessive moisture condition in the root zone that might reduce oxygen
availability (Savvas, 2009). Poor oxygen diffusion leads to lack of oxygen which causes die
off roots (Verhagen, 2009).
In addition, peat breakdown could lead to a more open structure with higher air filled
porosity. Weakly decomposed peat has an open structure which is sensitive to breakdown. In
practice is seen that peat materials with a high bulk density are less sensitive than those with
low bulk densities. Cattivello et al (1997) reported that particle size it self is not an important
factor in shrinkage of peat materials. On the other side, Verdonck and Demeyer (2004)
revealed that particle size is an important factor in determine the water and air relationship in
peat substrate.
These interactions show, that physical and biological proprieties of growing media
cannot be considered independent from cultural measurers directly (e.g. fertilization) or
indirectly (e.g. affecting carbon partitioning with the plant) the plant nutritional situation in
the root zone. In this context it has to be pointed out that microbial communities associated
with roots and growing media are extremely complex.
The most common growing media for greenhouse vegetable production are rockwool
(Tayson et al, 2001), perlite and gravel (Hamdy et al, 2002) as mineral medias. Currently,
more than 50 % of greenhouse vegetables are produced using rockwool as a growing media
(Zheng et al, 2009).
However, several factors must be considered when selecting a media for soilless
hydroponic vegetable production. A media must drain freely, especially as it ages, be easily
aerated, have good water holding capacity, be nontoxic, and free of weeds, pests, disease and
chemicals (Cantliffe et al, 2003). Furthermore, the media must be inexpensive (Cantliffe et al,
2001).
2.1.1. Rockwool
Rockwool is the most widely used inert substrate in soilless systems (Martyn, 1996;
Bohme et al, 2006) and it has been considered the most suitable medium to utilize the
following hydroponics-characteristics:
(ii) May hold a large amount of solution and has optimal aeration
Rockwool 80 18 7
*
Prasad (1997)
On contrast, it can’t be denied that using of rockwool slabs has also many
disadvantages such as:
(i) There is a substantial loss of unused water and nutrient elements when the nutrient
solution flows from the slabs
(ii) Root growth may become as massive as to fill the slab leading to root death (O 2
starvation) and root may grow out of the drain holes in the base of the sleeve
enclosing the slab
(iii)The slab requires periodic leaching to remove accumulated salts is order to prevent
a soluble salt buildup, a determination based an electrical conductivity (EC)
measurement made on a draw liquid sample from the slabs or effluent from the
slabs (Benton, 2005).
Fig. 2. Porosity of inorganic materials (a) volcanic scoria, (b) perlite and (c) pumice.
(Anicua-Sanchez et al, 2008)
(a) (b)
(c)
2.1.2. Peatmoss
Some growers use non-mineral media such as pine bark, coconut coir and sphagnum
peat mixtures because of their satisfactory aeration and water holding capacity (Cantliffe et al,
2003; Iapichino e al, 2007).
Peat has been the main raw material for industrially produced growing media, in
particular in greenhouse production (Sarkka et al, 2004). The availability of peat in large
quantities of relatively high and homogenous quality is necessary for different purpose. The
high percentage of peat in substrates makes the quality aspects very important (Keijzer, 2003).
*Cantliffe et al (2003)
Fig. 3. Porosity of organic materials (a) coconut fiber, (b) sphagnum peat and (c)
vermicompost. (Anicua-Sanchez et al, 2008)
(a) (b)
(c)
Table 4. Physical properties of organic media used in greenhouse culture*
In this respect, Verdonck and Demeyer (2004) declared that the physical of
growing media become more and more important for optimal plant growth. As they can
not be changed during the growing period, these properties should be optimal from the
beginning. In contrast, the chemical properties can be adapted to optimal conditions
during the growth
Physical property
Media type Moisture Air porosity (%) pH
capacity (% vol.)
Coir 33 16 5
Physical property
Media type Volume Volume
+
% air % EWA
Peat 0-1 mm 6.9 35.5
Light sphagnum peat possesses a range of positive and characteristics, including low
bulk density, good water holding capacity.
On the other side, the difficulty of obtaining positive impacts of AM on plant growth
in peat based growing media has been recognized for some time (Estaun et al, 1994).
There are several possibilities for this like the excess amount of P in the substrate, the
low anion holding capacities of sphagnum peat lead to increase the need for frequent fertilizer
application in greenhouse culture, and AM suppressive microorganisms in peat may be
another possible reason for weak AM symbiosis in peat based growing media. Also, peat
growing media have high populations of fungi and bacteria, in particular actinomycetes,
which may affect symbiosis efficiency of AM (Vestberg and Kukkonen, 2009). In addition,
extra cellular enzymes like ß-glycosidase, phosphat ase, sulphatase, xylosidase, esterase
present in peat (Freeman et al, 1995) may interact with AM.
Linderman and Davis (2003) demonstrated that different peat substrates indeed have
some deleterious effect on mycorrhiza formation and the potential for mycorrhiza induced
plant growth enhancement under P limiting conditions.
The authors concluded that the nature of the inhibitory factor, however, remains
unknown. This factor may be found in the organic chemistry of peat. Peat consists of mainly
humines, humic and folic acids, cellulose and hemi cellulose, bitumen, pectin and small
amount of mineral substances. Some of this component may interact with AM.
However, this effect can vary also with AM strain (Corkidi et al, 2004). In contrast,
humic acids have shown a positive impact on AM development and function (Gryndler et al,
2005).
Certain peat qualities can even possess properties that seem to have a negative effect
on AM symbiosis. Incompatibility between AM and peat, the reasons of which are still
largely unknown, is maybe the largest obstacle for using AM in peat-based growing media
(Vestberg et al, 2000; Linderman and Davis, 2003).
Slabs of peat and coconut fibers have been used successfully and with increasing a
mount (Bohme and Vorwerk, 2003). Tahvonen and Kemppainen (2008) reported that light
sphagnum peat has proved to be the best and most reliable substrates for plants in
greenhouses when aiming for higher yields.
So that, the effect of different substrates will be reviewed as follows in the next
chapters:
On the other hand, Cantliffe et al (2003) declared that type of media (perlite, coarse
perlite, and pine bark) did not affect stem length in pepper plants. Similarly, Shinohara et al
(1999) found no differences in growth characters between tomato plants grown in coconut
fiber or rockwool.
In cucumber, number of leaves, stem diameter as well as plant height in plants grown
in pumice or perlite did show any differences among the substrates (Abul-Soud et al, 2003).
Recently, AL-Rawahy et al (2009) did not find differences between the two substrates
(wood straw and date palm leaf straw) in terms of plant height and stem thickness for
cucumber plants grown in open soilless system. Also, Parks et al (2009a) revealed no
difference between medium contained: equal share of sand, perlit, composted pine bark and
coir, by volume, when compared to another medium contained 25 % sand, 25 % perlite, 50 %
composted pine bark in terms of cucumber shoot dry weights.
However, Inden and Torres (2004) declared that fresh and dry weights of tomato were
significantly higher in plants grown in perlite plus carbonized rice hull, while no differences
th
in fresh and dry weights of the 4 leaf were recorded between plants grown in rockwool or
coconut coir substrates. In addition, Ikiz et al (2009) studded the effect of three different
application treatments of mycorrhiza on pepper plant grown in soilless culture. The authors
found that dry weight of the plants grown in perlite medium had 28 % increases by one time
mycorrhiza application and 41 % increases by two time mycorrhiza application if compared
to control.
However, effects of different substrates on crop yield and its component are varying
among grown crops. In this respect, Maniutiu et al (1995) found that cucumbers plants grown
in bags of peat showed significant increase in early and total yield compared to the plants
grown in straw beds.
Moreover, Lee BeomSean et al (1999) pointed out that greater marketable yield and a
lower incidence of abnormal fruits were observed on a substrate of perlite with rice hull or
with carbonized rice hull as compared to the pure perlite substrate. In addition, Broeck (1999),
declared that cucumber plants cultivated in pumice produced the highest fruit number, with
slightly lower average crop yield than rockwool substrate.
Mohamed and Enzy (2001) found that early yield of sweet pepper plants was
increased significantly when plants grown on sandy soil, sand bags and gravel systems when
compared with NFT system. Moreover, Tuzel et al (2001) mentioned that tomato grown in
perlite + peat resulted higher fruit number per plant than plants grown in perlite alone.
In connection, Colla et al (2003) reported that marketable yield per plant was
significantly higher in cucumber plants grown in perlite and coir than those grown in
rockwool. The authors also mentioned that high fruit number and average fruit weight of
plants grown in perlite and coir caused these significant difference compared to plants grown
in rockwool. These results were confirmed by Parra et al (2009) on tomato, which found that
marketable yield for tomato grown in coir was significantly higher than those grown in
rockwool under open system condition.
Moreover, Peyvast et al (2008) pointed out that total marketable yield (9.0 and 7.9
-1 2
kg.m ) and numbers of cucumber fruits (102 and 92 m ) were significantly higher with plants
grown in peat than perlite, respectively. The authors found that peat substrate produced the
2
highest fruit fresh weight with 9.8 kg.m followed by perlite and perlite + rice hull with an
2
amount of 8.6 and 8.1 kg.m , respectively. Meanwhile, different substrates had no significant
effect on the non-marketable yield. Similar trend was obtained by Show et al (2004) on ‘Beit
Alpha’ cucumbers.
Recently, Parks et al (2009 b) mentioned that mean fruit weight was approximately
48 % higher for cucumber and 12 % higher for tomato grown in coir compared to the other
substrates (three component including sugar cane waste, peanut husks, soybean, peat or
volcanic rock).
On the other hand, the highest tomato fruit yield was obtained for plants grown in
rockwool, compared with plants grown in expanded clay or polyurethane foam (Chohura and
Komosa, 2000) or coconut fibers (Benko et al, 2008). In addition, Bohme et al (2008) found
that yield was very low in cucumber plants grown in peat slabs compared to those grown in
rockwool. The authors explained this increase to the rockwool ability to absorb more nutrients
than peat slabs.
Meanwhile, many authors found that yield of cherry tomato cultivated on coconut
fiber was similar to the yield on rockwool (Holtmann and Kobryn, 2003; Inden and Torres,
2004). Lopez-Galarza et al (2005) pointed out that type of substrate (coir dust, perlite and
rockwool) did not affect yield of watermelon plants
In this respect, Cantliffe et al (2003) declared that type of media (medium perlite,
coarse perlite, and pine bark) had no significant effect on fruit yield of cv. Beit Alpha
cucumbers (Show et al, 2004) as well as fruit set and total number of nodes per plant in
peppers. While, in strawberry, plants grown in perlite produced a higher marketable fruit
number and fruit weight per plant compared to pine bark and mixture of 2 peat: 1 perlite.
Also, Schahinrokhsar and Tavusi (2008) found no significant differences in yield, fruit
number and fruit weight of strawberry grown in coco peat or bark substrates.
In cucumber, Schroeder and Sell (2009) found no significant differences in yield for
‘Euphoria’ and ‘Aviance’ cultivars grown in peat or composted substrate.
Recently, no difference in fruit number and fresh weight of cucumbers and tomatoes
were found between medium contained 25 % sand, 25 % perlite, 25 % composted pine bark
and 25 % coir by volume when compared to medium contained 25 % sand, 25% perlit, 50%
composted pine bark (Parks et al, 2009 a).
Tuzel et al (2001) found that fruit of tomato showed significant increases in dry
matter, vitamin C and total soluble solid content when grown in volcanic ash compared to
perlite substrates. Also, fruits of cherry tomato grown on coco-fiber contained much more
total sugars, while those grown on rockwool decidedly in vitamin C (Holtmann and Kobryn
2003). Similarly, Inden and Torres (2004) found that the high soluble solids content was
higher in plants grown in coconut coir (7.3) followed by rockwool (6.6 O Brix) and perlite
(5.6O Brix).
In addition, Parks et al, (2009 b) studied the effect of four substrates on quality of
cucumber fruits. The authors found that nitrate concentration in cucumber was particularly
higher for sugar can waste, peanut husks and peat medium, while vitamin C recorded higher
content in the coir medium.
On the other hand, Guler et al (1995) did not find differences in soluble solid content
among melon fruits grown in rockwool, perlite, coir slabs or coir dust + perlite mix. Also,
quality of cucumbers grown in straw beds or bags of peat substrates (Maniutiu et al, 1995) did
not show any differences among substrates. Similar results were obtained with tomato grown
in coconut fiber and rockwool (Shinohara et al, 1999).
Also, strawberry plants grown in coco peat and bark substrates did not reveal
significant differences between the two substrates (Schahinrokhsar and Tavusi, 2008). Al-
Ajmi et al (2009) found that K and P concentrations were higher in fruits of cherry tomato
grown in zeolite if compared to perlite substrate, while fibbers and protein contents were not
affected by substrates.
reaction for the fixation of nitrogen (N + 3H2 + Energy -> 2NH3) is identical for both the
chemical and the biological processes. Living organisms use energy derived from the
(ii) Some have adapted to form symbioses; others have intimate entophytic
associations with plants.
(iii)Others live in close association in the plant root zone (rizosphere) without forming
intimate endophytic symbioses (Rai, 2005).
Azotobacter is Gram negative bacteria, polymorphic i.e. they are of different sizes and
shapes. Their size ranges from 2-10x1-2.5 µm. Young cell possess peritrichous flegella and
are used as locomotive organs (Fig.4). They also produce polysacharides. It is sensitive to
o
acidic pH, high salts, and temperature above 35 C. (www. 2).
*(www. A)
Nitrogen fixing bacteria obtains the required energy from the oxidation of
carbohydrates which have been formed by the photosynthetic activity of green plants. The
energy for photosynthesis comes from sunlight.
The energy for bacteria is therefore indirectly derived from a universally available and
inexhaustible source. The direct source of energy (carbohydrate) for bacteria is therefore
potentially available wherever conditions permit the growth of autotrophic organisms.
However, the ability of a biological system to fix nitrogen is dependent on the presence of a
particular enzyme “nitrogenase”. This enzyme cataly zes the conversion of N 2 into a reduced
form (ammonia combined with certain organic compounds) which can then be used for
growth by microorganisms and higher life forms.
*(www. B)
The nitrogenase system consists of two different protein molecules (enzymes) which
must function together in the nitrogen fixing process. The ammonia resulting from fixation is
rapidly incorporated into certain amino acids, such as glutamine or alanine. The nitrogen may
then be transferred to other amino acids and nitrogen-containing compounds by a variety of
commonly occurring amino transfer reactions (Hubbell and Gerald, 2003).
Azospirillum sp. are known mainly for their ability to produce plant hormones as well
as polyamines and amino acids in culture (Thuler et al, 2003), but they are also involved in
the biological fixation of nitrogen and the increased activity of glutamate dehydrogenase and
glutamine synthetase (Ribaudo et al, 2001).
Table Sea 35 6.
Total 135
12
Type of fixation N2 fixed (10 g per year)
Non-biological
Industrial 50
Combustion 20
Lightning 10
Total 80
Biological
Agricultural land 90
Forest and non-agricultural land 50
Estimates of the amount of nitrogen fixed on a global scale*
*(Bezdicek and Kennedy, 1998)
3.2. Arbuscular Mycorrhiza
Mycorrhiza is involved in the most durable, intimate and important symbiosis on the
earth (Allen, 1996; Hodge, 2000), and form mutualistic symbioses with a vast majority of land
plants (Smith and Read, 1997). Possibly more than 80 % of all terestric plants form
mycorhizal symbioses.
Mycorrhiza is traditionally classified into two types, ectotrophic and endotrophic. This
classification was based on the position of the fungal hyphae in relation to the root tissues of
the plant (Raj, 2005). Ectomycorrhizal hyphae ensheath the root cells, but do not penetrate
them, i.e. they remain extra cellular. While, endomycorrhizal hyphae are intracellular as they
actually penetrate the root cells of plants.
This classification is now regarded as too simplistic, and has been superseded by a
nomenclature identifying 7 mycorrhizal types. The term ectomycorrhiza remains, as the fungi
within this group are reasonably homogenous, but a sub-group, ectendomycorrhizas, is added.
Within the endotrophic group, 5 distinct types have been classified as follows:
The association between plants and their root colonizing mycoorhiza fungi is a
functional symbiosis in which the mycorhizal fungus is obligatory or facultative dependent on
host photosynthesis and energy. The symbiosis develops in the roots where the fungus
colonizes the cortex and obtains carbon from the plant while facilitating the transfer of
mineral nutrients to the root cells (Harrison, 2005).
The most important benefit of mycorrhiza is the increase in nutrient absorption, such
- ++ ++ +
as H2PO4 , Zn , Cu and NH4 . These benefits lead to improve the nutritional status of the
plants, to widen their ability to adapt to different ecosystems, and enhance the tolerance to
stressful biological and non-biological factors (Siqueira and Saggin Junior, 1995).
Thus, resulting from the more nutrient uptake, plant growth is generally improved
when mycorrhizal fungi colonize the root system (Rai, 2005). Moreover, when there is a
water deficit, the direct transportation of water to the roots can be interrupted and, in this case,
the external hyphae also acquire an important role in the absorption of water (Allen and Allen,
1986).
*(www. C)
*(www. D)
Besides the nutritional benefits, numerous non-nutritional benefits have been
attributed to the AM, including:
absorption of CO2, photosynthesis (Mathus and Vyas, 1995; Ruiz-Lozano et al, 1995;
Goicoecheta et al, 1997; Rilling et al, 2002).
All organic growing media contain microorganisms (Carlile and Hammonds, 2008).
The species and number of these microorganisms depends on several factors including the
constituents of media, their associated moisture and nutrient content, pH, C:N ratio and types
of carbon. Raw peat has a low microbial content, linked to its low pH and high lignin content
(Kavanagh and Herlihy, 1975).
A shift of perspective occurred when Carlile and Wilson (1991) presented their review
on microbial activity in growing media which highlighted the influence of microorganisms
associated with growing media regarding their physical and chemical qualities. They
concluded that this manipulation of microbial growth in soilless culture may offer
considerable benefits to growers and may form an integral part of commercial use of soilless
media in a near future (Al-Moshileh, 2007).
Moreover, inert growth systems such as rockwool are deficient in microbial activity,
and applying mycorrhiza inocula to these systems may not only result in microbial
mycorrhiza formation, but also in introduction of mycorrhiza associated bacteria (Mansfeld-
Giese et al, 2002). In this respect, Bin et al (2008) found that mycorrhiza associated bacteria
from the genus Paenibacillus have been shown to promote growth of cucumber plants.
In comparison among different N-Fixing bacteria, Gupta et al (1995) pointed out that
inoculation tomato plants with Azotobacter was most effective in increasing the total dry
weight, root and shoot length, followed by Azospirillum.
Indeed, several studies indicated that Azospirillum can increase the growth of various
crops. These include carrot, eggplant, pepper, oak, sugar beet, sugarcane and tomato (Bashan
et al, 1989; Bashan and Holguin, 1997).
In this respect, Barakat and Gabr (1998), under inoculation with Azospirillum,
recorded a significant increase in the tomato seedling height, leaf area, fresh and dry weight,
but not in number of leaves per seedlings. In accordance, Cho Ja Yong et al (1998) mentioned
that Azospirillum sp. promoted early growth of cucumber and tomato plug seedlings.
These results are in harmony with Tantawy (2000), who found that inoculation of
tomato seeds with Azospirillum increased stem length, plant dry weight and plant fresh
weight, which reached 13.5, 35 and 52 %, respectively. In connection, Garib (2002), found
that inoculation of cucumber with Azotobacter and/or Azospirillum promoted different growth
characters i.e. number of branches, number of leaves and leaf area under greenhouse
condition.
These positive effects resulted from bacteria inoculations are related to their ability to
improve plant nutrition by fixing atmospheric N, and influence plant growth by production of
plant enzymes (Gruda et al, 2004), hormones such as gibberellins (Gutierez-Manero et al,
2001) and cytokinins (Timmusk et al, 1999).
On the other hand, Gomma (1995) revealed no response of tomato seedlings, as plant
height, fresh and dry weight of plants, if inoculated by Azotobacter and Azospirillum. Similar
trend was mentioned by Gruda et al (2008), whom observed that additives of Bacillus subtilis
did not affect fresh and dry weight, number of leaves as well as leaf area of primula and
begonia plants.
However, it is difficult to explain the role of Bacillus subtilis since a little has been
published on the physiological mechanisms involved as opposed to the agronomic effect of
this microorganism (Saleh et al, 2005).
On the other hand, Douds and Reider (2003) found that inoculation of sweet pepper
seedlings with G. intraradices prior to transplanting decreased relative seedling growth to
control. Sabin and Larsen (2005) did not reveal differences between mycorrhizal and non-
mycorrhizal tomato plants in terms of growth parameters.
Moreover, Yesilova and Karaca (2007) mentioned that inoculation of melon plants
with G. entunicatum increased significantly plant and root lengths as well as shoots dry
weight when compared with control plants under the laboratory condition, while plant growth
parameters were not significantly differ under greenhouse conditions. Recently, application of
mycorrhiza to pepper plant improved plant growth than control under soilless culture
condition (Ikiz et al, 2009).
3.4. Yield and its component
In two decades of field experiments, general consensus is that in 60 to 70 % of the
cases Azospirillum application results in a significant crop yield increase (Okon and
Labandrea-Gonzalez, 1994).
In this respect, Miles (2007) mentioned that Azotobacter increase plant yield primarily
by fixing molecular nitrogen, but it is also reported to synthesize auxins, vitamins, growth
substances and antifungal antibiotics, which have beneficial effects of this bacterium on seed
germination. The author revealed that Azotobacter spp. inoculants can increase crop yields on
barely, cabbage, maize, oat, onion, potato, rice, sugarcane, tomato, wheat by 7-12 %.
Regarding to the effect of arbuscular mycorrhizal (AM) fungi inoculation, Douds and
Reider (2003) found that inoculation of sweet pepper seedlings with Glomus mosseae prior to
transplanting showed significant increase in fruit yield, while Glomus intraradices decreased
yield relative to control.
In this respect, Sabin and Larsen (2005) studied the possibility of implementing AM
fungi in commercials cucumber production. The authors found, in the first production cycle
from January to July in two greenhouses, that AM did not influence cucumber yield, but in the
third nursery inoculation with AM resulted in approximately 4 % more in first class cucumber
compared with plants without mycorrhiza, which would correspond to an extra profit of
approximately 50,000 Euro/ha in the spring production cycle.
On the other hand, Gul et al (2007) documented that inoculation of tomato with
Bacillus subtilis or Trichoderma horyianum did not affect vitamin C or fruit quality
characteristics.
In connection, Demir (2004) mentioned that leaves of pepper plants inoculated with
Glomus intraradices showed better phosphorus, dry matter, chlorophyll and amount of some
reducing sugars (fructose, alpha glucose and beta glucose) comparing to un-inoculated plants.
These results are in harmony with Neumann (2005) who reveled that AM fungal inoculation
increased phosphorus uptake of tomato cv. Golden Queen.
In cucumber, nitrogen, chlorophyll and dry matter content in leaves were increased
under greenhouse conditions using combination of Azospirillum + Azotobacter + phosphate
dissolving bacteria (Garib, 2002).
In addition, in two pot experiments, cucumber (Cucumis sativus L. cv. Jinlu No. 3)
seedlings were each inoculated with one of three arbuscular mycorrhizal fungi, Glomus
mosseae, Glomus intraradices and Glomus versiforme. Contents of nitrogen and phosphorus
in roots and magnesium, copper, and zinc in shoots were increased by inoculating treatments,
while potassium and iron contents in shoots decreased significantly (Changxian Wang et al,
2006).
o Passandra F1
Introduced by Enza-Zaden company, Holland. It has good growing vigorous open plants,
giving side-shoots, 2-3 fruits/node and top yielding. The plants and fruits are bitter free. The
fruits are dark green, slightly ribbed and cylindrical with a length of 150 mm “mini fruit
type”. It is powdery mildew and scab tolerant as we ll as recommended for autumn and spring
crops.
o Girola F1
Introduced by Enza-Zaden company; Holland. It has strong vigorous plants, produces
high early and total yield. The fruits are dark green, smooth, slightly ribbed with a length of
330 mm “long fruit type”. It is leaf-spot, scab res istance and recommended for autumn as
well as winter planting.
Microorganisms inoculation
Nitrogen fixing bacteria Aztobacter chroococcum CCM 1921 and Azospirillum
brasilense CCM 3862 were obtained from Czech Collection of Microorganisms (CCM)
institute, Masaryk University, Faculty of Science, Brno.
Bacteria were applied to transplants, grown in 80 mm trays filled with sphagnum peat as
a transplants growing media, using liquid culture from each strain at rate of 1 ml liquid
9
culture per plant (1 ml contains 10 cells of bacteria) as described by Valentine et al (2002).
Arbuscular mycorrhiza fungi Glomus mosseae were mixed to sphagnum peat as a
growing media at rate of 3 g per plant as recommended by Symbio-m Company (CZ).
Microorganisms applications were replicated 2 times during transplants growing stage.
th th
The first inoculation dose was applied around root zone at 14 of March 2006 and 26 of
February 2007, respectively. Second inoculation dose was applied 15 days after the first
treatment.
The inoculation treatments and its mixtures were applied as follows:
1. N1 (Azotobacter chroococcum)
2. N2 (Azospirillum brasilense)
3. G (Glomus mosseae)
Training system
The basic principles of training the plants to uniformities distribute the foliage
throughout the entire greenhouse so that the leaves can intercept maximum light.
Single crop wires was placed 2.5 m above the ground over each row of plants. The
strings for plants were then tied alternately to the overhead wires so that the plants were
inclined away from the row on each side. The plants let to grow in a V- arrangement down the
row, allowing fruit to hang away from the main stem.
Pruning
The pruning method has been prune to result in early and high fruit production as
described by Vogel et al (1996) as following:
1. Plants were trained to be overhead wire and the growing point was removed tell
500 mm for cultivar ‘Passandra’, as a mini fruit ty pe, and 700 mm for
cultivar ‘Girola’, as a long fruit type.
2. A small loop of string was tied around the wire and the main stem below the top
leaf to prevent the plant from slipping down the string.
3. All side shoots were cut after producing one fruits at a very early stage, except
one fruit - one leaf was left for cultivar Girola. While with cultivar Passandra, it was
allowed to produce one more fruit and one more lateral leaf after the first fruit has
been trained.
4. Three or four leaves have been selected when the main stem reached the top of
wire, then other 2 branches were allowed to grow down. Only the laterals were
topped when necessary (when they touch the ground).
Data recorded
I. Vegetative growth characteristics
Three vegetative growth plant samples were collected randomly from each replicate after
1. Plant height.
2. Numbers of leaves/plant.
3. Total leaf area/plant.
4. Fresh and dry weights of leaves.
II. Yield
th th th
Harvest, at marketable green mature stage, started on 4 and 15 April and finished on 18
th
and 5 June 2006 and 2007, respectively. The following yield parameters were recorded:
-2
1. Early yield, the sum of the first five pickings (kg.m ).
-2
2. Total yield, the all harvesting pickings (kg.m ).
3.
III. Fruit quality
To estimate fruit quality, samples of five fruits at the marketable green stage of the tenth
and ninths picking of the first and second season, respectively were chosen randomly from
A. Physical properties
1- Fruit length and diameter were estimated by Vernier Caliper instrument in mm.
2- Average weight of fruit (g).
3- Quality according to the EU norm No. 46 3155.
B. Chemical properties
decrease in O2 levels in rooting media which reflected on root activity, nutrient uptake
absorption and growth vigor (Holtman et al 2005).
In this respect, it can be suggested that these obtained differences in growth
characteristics of cucumber plants could be related to distinction in chemical and physical
properties of rockwool and peatmoss.
On the other hand, plant height did not record significant differences between the used
substrates (rockwool or peatmoss) in both growing seasons. Similar results were obtained by
Cantliffe et al (2003) who pointed out that type of growing media (perlite or pine bark) did not
affect stem length in pepper plants.
In connection, Abul-Soud et al (2003) found that plant height of cucumber plants
grown in pumice or perlite did not show any differences among the substrates. Meanwhile,
AL-Rawahy et al (2009) mentioned that plant height and stem thickness for cucumber plants
grown upon wood straw and date palm leaf straw in open soilless system did not show
differences between the two substrates.
In this respect, it can be suggested that non-significant differences might be not related
to the type of used substrates, but might be due to other factors, i.e. genotype. (Hanafy Ahmed
et al 2004; Abdelaziz et al, 2007).
1.3. Effect of microorganisms inoculation
Table (8) clears that inoculation of cucumber plants by N-Fixing bacteria and/or
Glomus did not affect vegetative growth characteristic in both growing seasons if compared to
control treatment, with some exceptions.
50
40
30
20
10
2 G 1 G+N2 2 l
o
N1 N N1+N2 N r
+N n
t
G+ o
G+N1 C
4,0
3,5
g/leaf
3,0
2,5
2,0
1 2 2 N1 N2 2 l
o
N N G
+N + + +N ntr
1 G G 1 o
C
N G +N
2,2
2,0
m
1,8
1,6
1,4
N1 2 2 G 1 2 G+N1+N2 ontrol
N N G+N G+N
+
N1 C
Rockwool Peatmoss
Similarly, plants grown in peatmoss and inoculated with Azospirillum showed higher
value of leaf fresh weight (31 g), when compared to other treatments in the first season (Fig.
13). No differences were found in the second season (Table 16).
While in Table (18), cucumber plants grown in peatmoss and inoculated with (Glomus
mosseae + Azotobacter) led to a significant increase of leaf dry weight (4 g) in the first
season, as shown in Fig. (14).
Fig. 11. Effect of interaction between substrate x microorganisms inoculation
on leaf area of cucumbers in 2006
1200
900
2
cm
600
300
N1 N2 G G+N1 G+N2 2 l
o
N1+N2 r
G+N1+N t
on
C
Rockwool Peatmoss
50
40
30
20
10
N N tro
G+N1+N n
G Co
Rockwool Peatmoss
40
30
g/leaf
20
10
N N N
+
1
N
Rockwool Peatmoss
5,0
4,0
3,0
g/leaf
2,0
1,0
N1 2 2 G G+N1 2 l
G+N1+N2 o
N 1+N +N t
r
G Con
N
Rockwool Peatmoss
20
16
-2
mKg.
12
8
2006 2007
Rockwool Peatmoss
L.S.D. NS NS NS 1,4
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
and antifungal antibiotics (Miles, 2007). Furthermore, presence of plant growth promoting
rhizobacteria (PGPR) might not result in signs of visible plant growth but contribute to plant
yield by reducing or suppressing disease incidents. It may be more practical to say that PGPR
help plant either directly via visible growth promotion or indirectly via disease control (Rai,
2005).
In this respect, it can be suggested that microorganisms inoculation did not play a vital
role to suppress disease function in the first season. However, it can be said that dual
inoculation of N-Fixers was partly more active in control disease and improve yield
productivity in the second season.
2.1.4. Effect of interactions
Inoculcated plants of cultivar Passandra with Glomus recorded the highest significant
-2
values of early yield (0.9 kg.m ) if compared to other treatments in the first season only as
shown in Table (21). Other interaction treatments did not show significant differences in both
seasons, Tables (21-22).
In terms of total yield, cultivar Passandra grown in peatmoss revealed higher
-2 -2
significant increase (17.5 kg.m ) in the first season, and 13.6 kg.m in the second season
when grown in rockwool, as shown in Table (19).
Moreover, total yield of cultivar Passandra presented better significant increase when
-2
inoculated with Glomus (18.7 kg.m ) followed by those inoculated with Azotobacter (18.2
-2
kg.m ) in the first season (Fig. 16).
20,0
17,5
-2
15,0
Kg.m
12,5
10,0
N1 N2 2 G 1 N2 l
o
N N1+N2
1+N + + ntr
o
N G G C
+
G
N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae
20,0
17,5
-2
Kg.m
15,0
12,5
10,0
N 1+
N1+ Cont
N
+
G
N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae
260
250
240
g
230
220
Con
However, these values did not reach to significant level but pointed out that
microorganisms inoculation had positive effects on increase cucumbers fruit weight.
Same results were recorded by Shahaby et al (1993) and Miles (2007) on tomato, Li
Ning et al (1997) on pumpkin, Omar and El-Kattan (2003) on cucumber whom found that
inoculation vegetables crops with microorganisms enhanced yield and its component.
In this respect, it can be suggested that preplanting microorganisms inoculation
treatments had favorable influence on average fruit weight. This might be a result of N-Fixers
effects on release more available nitrogen in root zone.
In accordance, improved levels of nitrogen and other nutrients led to positive effects
on many vital processes, i.e. chlorophyll, enzymes, photosynthesis and endogenous hormones
synthesis which consequently affect growth and yield (Marschner, 1995, Hanafy Ahmed,
1997 and Garib 2002).
In general, combination of N-Fixers or the single inoculation of Glomus enhanced the
vital process of plant organs and caused that increase in average fruit weight of cucumbers.
However, these effects of microorganisms reflected positively on average fruit weight than
fruit length, diameter or number. In particular, physical component such as fruit length and
diameter are more correlated to geno-type and less responding to microorganisms inoculation.
40
30
20
10
Passandra Girola
Rockwool Peatmoss
Meanwhile, both cucumber cultivars produced higher values of fruit weight when
grown under rockwool than peatmoss in the second season (Fig. 20).
Moreover, the interaction between cultivar x microorganisms inoculation or between
substrates x microorganisms inoculation did not affect fruit length, diameter and weight in
both growing seasons (Tables 27 to 30). The only exception found in fruit diameter of cultivar
Girola which recorded the highest significant increase (40 mm) with Azotobacter+
Azospirillum + Glomus inoculation if compared to the inoculation treatments (Tables 33).
Fig.20. Effect of interaction between cultivar x substrate on
cucumbers fruit weight in 2007
400
300
200
g
100
0
Passandra Girola
Rockwool Peatmoss
In terms of number of fruits per plant (Table 31), it can be observed that cultivar
Girola revealed higher significant increase (22) when inoculated with Glomus + Azotobacter
+ Azospirrillum than control (20.6). While, cultivar Passandra trended similarly and produced
significant increase of number of fruits per plant when inoculated with Glomus (41), if
compared to other treatments in the first season, as shown in Fig. (21).
50
40
30
20 c
10
1 2 N2 G 1 2 +N2 l
N N tro
+ G+N G+N n
1 o
N N1 C
+
Passandra Giola G
40
35
30
25
20
75 Extra
50 Class A
Class B
25 Unmarketable
0
100
75
Extra
50
Class A
25 Class B
Unmarketable
0
Season 2006
N1 11,4 4,23 1,04 0,17
N2 10,7 4,16 1,73 0,20
N1+N2 13,1 2,53 0,75 0,00
G 12,1 3,18 1,29 0,02
G+N1 11,8 3,67 1,09 0,23
G+N2 11,4 3,67 1,29 0,15
G+N1+N2 10,8 3,01 1,49 0,31
Control 10,5 3,76 1,19 0,02
L.S.D. 1,47 0,94 0,44 NS
Season 2007
N1 7,8 0,88 0,28 0,03
N2 6,9 0,81 0,44 0,20
N1+N2 8,1 1,34 0,25 0,27
G 8,1 0,64 0,51 0,15
G+N1 7,4 0,96 0,51 0,05
G+N2 8,1 0,83 0,36 0,06
G+N1+N2 6,9 0,89 0,59 0,11
Control 6,9 1,00 0,24 0,20
L.S.D. NS NS NS NS
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
This might be attributed to the beneficial effect of microorganism’s inoculation to
increase the uptake of available nutrient from N and P in root zone of cucumber plants, and
then express their effects on flowering and fruit setting.
In this respect, it can be concluded that marketable yield was positively affected by
microorganisms inoculation. Similar conclusion was obtained by Veeranna et al. (2000) and
Hanafy Ahmed (2004).
16
12
-2
Kg.m
4
0
16
12
-2
Kg.m
+
G
These finding are confirmed in Tables (39-40). Data in those tables indicate that
inoculation of cucumbers with mixture of N-Fixers (Azotobacter + Azospirillum) produced
the lowest fruit yield of "Class A" in the first season. Moreover, no significant differences
were observed in terms of "Class B" or unmarketable fruits in both growing seasons (Tables
41-44).
In addition, higher significant increase in marketable fruit yield was found with
-2
cultivar Passandra (Fig. 25) when inoculated with Glomus (14.0 kg.m ) followed by
-2 -2
Azotobacter + Azospirillum (12.7 kg.m ) in the first season and 11.5 kg.m in the second
season. Meanwhile, in Table (38), cucumber plants grown in rockwool and inoculated with
mixture of N-Fixers (Azotobacter + Azospirillum) showed higher significant increase in
-2
marketable fruit yield (13.1 and 9.7 kg.m ) in both growing seasons, respectively (Fig. 26).
Moreover, cucumber plants grown in peatmoss and inoculated by Glomus +
-2 -2
Azotobacter (13.3 kg.m ) or Glomus + Azospirillum (13.2 kg.m ) revealed significant
-2
increase in the first season if compared to control (11.2 kg.m ), as shown in Fig. (27).
In the second season, Glomus inoculation treatment produced higher significant
-2 -2
increase of marketable fruit yield (7.2 kg.m ) if compared to control (5.8 kg.m ).
In general, it can be noticed that cucumber plants grown in rockwool found to produce
higher marketable fruit than those grown peatmoss, as shown in Table (38).
Fig.25. Effect of interaction between cultivar Passandra x microorganisms
inoculation on marketable fruit yield of cucumbers in 2006 and 2007
16
12
-2 Kg.m
8
4
N1
N1+N2 G G+N1 G+N2 G+N1+N2 Control
2006 2007
16
12
Kg.m -2
8
4
1 G+N2 l
1 2 2 G 1 o
N N N1+N N2
o
+N
G r
t
+ n
+N C
G
2006 2007
16
12
-2
Kg.m
N
+ +
N1 1 o
G+N C
2006 2007
Season 2007
L.S.D. NS 0,04 NS NS 12
4,0
3,8
g/100g dry mater
3,6
3,4
3,2
3,0
Rockwool Peatmoss
Passandra Girola
2250
fresh weight
2000
gmg.k -1
1750
1500
Rockwool Peatmoss
Passandra Girola
4,5
dry matter
4,3
4,0
g/100d
3,8
3,5
1 G G+N1 +N2 l
o
N N2 1+N2 N2 r
N G + Cont
G +N1
Passandra Girola
0,70
matter
0,60
0,50
g.100g dry
0,40
0,30
N +N G+N
N1 Cont
2006 2007
0,60
dry matter
0,50
g.100g
0,40
0,30
1 G +N1 G+N2 l
o
N2 N1+N2 +N2 r
N +N1 nt
G G Co
2006 2007
-1
Calcium in Table (53), showed significant increase (247 mg.kg ) in leaves of cultivar
Girola in the second season with Azotobacter inoculation treatment, while cultivar Passandra
-1
reveal the highest significant increase (263 mg.kg ) when inoculated with Glomus +
Azotobacter or Glomus + Azospirillum (Fig. 33).
300
fresh weight
250
200
mg.kg -1
150
N1 N2 N2 G G+N1 G+N2 N1+N2 rol
N1+ Cont
+
Passandra Girola G
Magnesium did not show significant differences with inoculation treatments in both
growing seasons if compared to control, as shown in Table (55).
As regard to the effect of interaction between substrate x microorganisms inoculation,
data in Table (48) show that nitrogen content in leaves of cucumbers recorded significant
increase (4.74 g/100g dry matter) when plants grown in rockwool and inoculated with
Azospirillum comparing to other treatments. In general, content of nitrogen were higher in
leaves of cucumber plants when grown in rockwool than peatmoss in the first season, while
no differences were observed in the second season.
Furthermore, phosphorus did not reveal significant differences between treatments in
both growing seasons Table (50).
In addition, potassium content in leaves of plants inoculated with Glomus recorded the
-1
best significant increase (1448 mg.kg ) when grown in peatmoss than rockwool (1398
-1
mg.kg ) in the first season only, as shown in Fig. (34).
Moreover, in the first season, calcium in leaves of cucumbers showed higher
-1
significant values (206, 205 and 204 mg.kg ) when plants grown in peatmoss and inoculated
with Azospirillum, Azotobacter or Glomus, respectively, as shown in Fig (35). In the second
season, contentss of calcium showed the best significant increase with Glomus + Azotobacter
-1
or Glomus alone (309 and 303 mg.kg ), respectively.
1500
fresh weight
1400
1300
-1
gmg.k
1200
1 2 2 G 1 G+N2 l
o
G+N1+N2
N N N1+N G+N ntr
o
C
Rockwool Peatmoss
400
fresh weight
300
200
-1
mg.kg
100
o
C
2006 2007
N P K Ca Mg
Treatments
g/100g dry matter -1
mg.kg fresh weight
Season 2006
Season 2007
3,5
3,0
g/100g
2,5
2,0
N1 N2 2
G G+N1 G+N2 2 rol
N1+N
+N on
t
G+N1
C
Passandra Girola
4,0
matter
3,5
g/100g dry
3,0
2,5
2,0
N
+ n
G N1+ Co
+
G
Passandra Girola
0,60
matter
0,50
0,40
g/100g dry
0,30
0,20
N1 2 N1+N2 G 1 G+N2 N2 Control
N N
G+ G+N1+
2006 2007
1200
weight
1100
1000
fresh
-1
mg.kg
900
800
N1 2 1+N2 G G+N1 +N2 2 Control
N G N1+N
N
+
Passandra Girola G
100
90
fresh weight
80
70
-1
mg.kg
60
N N N
+
G+ 1
N1 G+N
Passandra Girola
0,6
dry matter
0,5
0,4
g.100g
0,3
+
G +N1
Peatmoss Rockwool
The above results reveal that fruits of cultivar Girola showed better nutritional
contents than cultivar Passandra under the microorganisms inoculation treatments, as shown
in Figs (37, 38, 39 and 40). In addition, it can be observed that type of substrate, in general,
inhibited both of microorganisms and cultivars to express their interaction. In this respect, it
can be concluded that type of substrate is a critical factor affected nutritional status of
cucumber plants under the experiment conditions.
Fig.42. Effect of interaction between rockwool x microorganisms inoculation on
magnesium content in fruits of cucumbers in 2006.
120
weight
115
110
fresh
-1
105
mg.kg
100
N1 2 2 1 2 l
o
N G N N N2 tr
N1+N + + + n
1 o
G G N C
+
G
These could be related to the diurnal changes in pH or salinity of the nutrient solution
in growing media. In this respect, it can be suggested that many reasons could cause these
changes in composition of nutrient solution i.e., the low irradiance under greenhouse
condition reflected negatively on growth rate of greenhouse plants and leads to a lower
nutrient absorption and utilization. Also, due to low relative humidity under air-conditioning,
the relative rate water loss would normally be higher than the nutrient uptake, and this may
result in an overall increase in the nutrient concentration, thus creating salinity problem
(Awang et al, 2007).
IV. 4. Chemical quality
IV. 4.1. Vitamin C and total soluble solid (TSS)
4.1.1. Effect of cultivar
Data in Table (69) indicate that vitamin C and total soluble solid did not differ between
the two cucumber cultivars in both growing season, except cultivar Passandra which showed
significant increase in TSS (4.1 %) if compared to Girola cultivar (3.9 %) in the first season.
However, it can be noticed low variations between both cultivars in terms of vitamin C
and TSS values. In this respect, it can be concluded that type of cultivar did not affect vitamin
C or TSS contents. Similar suggestion was obtained by Hanafy Ahmed et al (2004) on sweet
pepper.
-1 % % -1 -1
mg.kg mg.kg mg.kg
Season 2006
Season 2007
160
140
fresh weight
mg.kg -1
120
100
N1 N2 +N2
G 1 G+N2 2 cont
G +N G+N1+N
1
N
2006 2007
4,50
4,25
4,00
%
3,75
3,50
1 2 l
o
N N2 N2 G G+N1 N2 N
+ + Contr
N1 G N1+
+
2006 2007 G
150
130
-1
mg.kg
110
90
N G+N
N
Passandra Girola
5,0
4,5
4,0
%
3,5
3,0
N1 N2 2 G 1 2 N2 ntrol
+
N G+N G+N +
N1 1 o
G+N C
160
140
-1
120
mg.kg
100
80
N1 N2 +N2 G 1 2 G+N1+N2 l
tro
G+N G+N Con
N1
Peatmoss Rockwool
Table (74) declares that total soluble solid presented the best significant increase (4.3
%) with cucumbers grown in peatmoss and inoculated with Glomus + Azotobacter +
Azospirillum, Fig (48). On the other hand, no differences in vitamin C and TSS were observed
in the second season.
These results show that mixture of Glomus and Azospirillum inoculation under
peatmoss substrate led to improve quality of cucumbers. These positive influences can be
explained according to the role of mycorrhizal fungi to cause useful physiological alterations
5,0
4,5
4,0
%
3,5
3,0
N1 2 +N2 G N1 G+N2 G+N1+N2 o trol
N
2006 G
N1 2007
n
+
C
In this respect, it can be concluded that type of cultivar or substrate, as single factor,
did not affect vitamin C or TSS content in fruits of cucumbers in both growing seasons with
some exceptions. In addition, inoculation of cucumber plants with Glomus + Azospirillum
improved vitamin C contents in fruits of both cucumbers cultivars, especially when
cucumbers grown in peatmoss substrate. Similarly, TSS contents were significantly higher in
fruits of cucumbers grown in peatmoss, but when inoculated with Glomus + Azotobacter +
Azospirillum.
IV. 4.2. Dry matter content, nitrate and total sugars concentrations
4.2.1. Effect of cultivar
Concerning the effect of cultivar on dry matter content, nitrate and total sugars
concentrations of cucumber fruits, data in Table (69) clear that dry matter of cultivar
Passandra fruits were significantly higher (2.3 and 3.1 %) than cultivar Girola (2.1 and 2.9 %)
in both seasons, respectively.
Total sugars showed similar trend in the second season with cultivar Passandra (19.6
-1 -1
mg.kg ) if compared to cultivar Girola (16.8 mg.kg ), while no difference were recorded in
the first season between both cultivars. On contrast, fruits of cultivar Girola presented higher
-1
significant increase of nitrate (145 and 351 mg.kg ) than cultivar Passandra (117 and 258
-1
mg.kg ) in the two growing seasons, respectively.
In this respect, it can be suggested that mini fruit type of cucumbers such as cultivar
Passandra, are more active to absorb more nutrients, hence rate of translocation and
accumulation of such nutrients in tissue of mini fruit type is higher than long fruit type, as
cultivar Girola.
accumulation was inversely related to accumulation of sugars. Similar findings were reported
by Blom-Zandstra and Lampe (1985) and Hanafy Ahmed (1996) on lettuce, Hanafy Ahmed et
al (1997) on Jew’s mallow and radish as well as Hanafy Ahmed et al (2002 a, b and c) on
rocket, lettuce and sweet pepper, respectively. In this respect, Blom-Zandstra et al (1988)
mentioned that the lettuce genotype which had higher nitrate concentration had also lower
concentration of sugars and organic compounds.
In addition, the increases in the concentration of total sugar may also be explained on
the assumption that, as already known, the nitrate accumulation in the vacuoles is not readily
available. However, when the nitrogen is not sufficient amount, the nitrate might remove from
the vacuoles to sustain the protein synthesis and the plant will store sugars and organic acids
to compensate for the declining osmotic value. This suggestion is in a good agreement with
that of Blom-Zandstra (1989).
30
25
-1
mg.kg
20
15
10
1 2 G G+N1 G+N2 l
o
N1+N2 G+N1+N2 r
N N nt
Co
Passandra Girola
In accordance, Vestberg and Kukkonen (2009) mentioned that peat growing media
have high populations of fungi and bacteria, in particular actinomycetes, which may affect
symbiosis efficiency of AM. In addition, extra cellular enzymes like ß-glucosidase,
phosphatase, sulphatase, xylosidase, esterase present in peat may negatively interact with
microorganisms population (Freeman et al, 1995).
V. SUMMARY
This experiment was carried out during the two successive seasons of 2006 and 2007
under the open soilless system in greenhouse at Faculty of Horticulture in Lednice, Mendel
University, Czech Republic. Transplants of cucumber (Cucumis sativus L. cultivars
‘Passandra’ and ‘Girola’) were arranged in 4 replic ates using split-split plot design. Each
cultivar was grown on rockwool and peatmoss as based growing media. Transplants were
inoculated by seven microorganisms treatments: N1 (Azotobacter chroococcum), N2
(Azospirillum brasilense), G (Glomus mosseae), and their combinations N1 + N2, G + N1, G
+ N2, G + N1 + N2 and control.
Generally, it can be observed that cultivar Passandra was exceeded cultivar Girola in
terms of plant height, number of leaves as well as leaf FW and leaf DW. On contrast, leaf area
nd
of cultivar Girola showed higher significant increase in the 2 season. Early and total yield of
both cultivars did not reveal much variation. These results indicate that each cultivar
expressed its effect on vegetative growth characteristics based on variation among both geno-
types. However, cultivar Girola presented higher significant increase in fruit length, diameter
and weight. Meanwhile, cultivar Passandra produced higher significant number of fruits/plant
in both seasons; combined with higher significant increase of grade “Extra” fruits were
nd
obtained in the 2 season.
Nitrogen, phosphorus and magnesium did not differ in leaves of both cultivars.
st nd
Moreover, K was significantly higher in leaves of cultivar Girola and Passandra in 1 and 2
seasons, respectively. Nitrogen in fruits of cultivar Passandra and P, K, Ca in fruits of cultivar
Girola revealed higher significant increases in the first season while N, P, and Mg did not
differ in leaves of the two cultivars, with some exceptions. Type of cultivar did not affect
vitamin C or TSS contents. Meanwhile, dry matter in fruits of cultivar Passandra were
st
significantly higher in both seasons; and total sugars in the 1 season. Nitrate, recorded higher
significant increase in fruits of cultivar Girola, but values still under the critical limits. In
general, it can be concluded that variations between cultivars are correlated to the gene-type,
and cultivars are varied in their yield productivity and quality.
According to the effect of substrate type on cucumber plants, data presented that plants
grown on peatmoss revealed significant increase in number of leaves/plant and leaf DW. Leaf
area and leaf FW showed different trends in both growing seasons. However, light significant
increase was obtained in early yield of plants grown on rockwool. On contrast, total yield of
st
cucumbers grown in peatmoss recorded higher significant increase in the 1 season. Similar
nd
trend was noticed with plants grown on rockwool in the 2 season.
Fruits of cucumbers grown in rockwool revealed lightly significant increase in fruit
length and weight. Meanwhile, cucumbers grown on peatmoss produced higher significant
st
fruits number/plant and marketable yield of grade “ Extra” in the 1 season and when grown
nd
on rockwool in the 2 season. Macronutrients in leaves and fruits as well as vitamin C, total
sugars and TSS in fruits of cucumbers did not differ between substrates, with some
exceptions. Fruits of cucumbers grown in rockwool showed significant increase in dry matter
- st
and NO3 in the 1 season. In this respect, it can be concluded that distinction in chemical and
physical properties between rockwool and peatmoss reflected on recorded results.
Cucumber plants inoculated by N-Fixing bacteria and/or Glomus did not affect growth,
early yield, fruit length, diameter and fruits number/plant in both seasons, with some
exceptions. However, it can’t be denied that microorganism inoculation treatments did light
positive effect on some of vegetative growth characteristics and yield component. Limited
light quantities inside greenhouse in winter season and/or the variation in pH and saltation of
the nutrient solution could inhibit the microorganisms formation under such conditions.
Furthermore, all inoculation treatment revealed better, but not significant, total yield.
st nd
These improve in total yield reached 8 and 21 % in the 1 and 2 seasons, respectively. In
addition, significant increase in average fruit weight was found when cucumber plants
st
inoculated with Azotobacter alone or combined with Azospirillum than control in the 1
season. Inoculating cucumber plants improved the marketable yield, contents of
macronutrients in leaves and fruits of both cultivars, with some exceptions.
Vitamin C in fruits of cucumber plants recorded the highest significant increase when
inoculated with Glomus + Azospirillum. In addition, total soluble solid revealed similar trend
st
in the 1 season when plants inoculated with Glomus + Azotobacter + Azospirillum.
Moreover, significant decrease in dry matter was observed with Azotobacter and Azospirillum
nd
inoculation if compared to control in the 2 season. Nitrate concentrations were significantly
increased with Azotobacter or Azospirillum treatments if compared to Glomus or control in
st
the 1 season.
Interactions between cultivars and substrate or microorganisms show preferable effect
on most of growth characteristics of cucumber plants. Particularly, N-Fixers alone/or
combined with Glomus improved growth of cucumber plants grown in peatmoss. Cucumber
plants grown in peatmoss and inoculated with Glomus + Azotobacter or Azospirillum, were
st
significantly higher in total yield in the 1 season than control. In general, it can be concluded
that growth and yield of cultivar Passandra grown in peatmoss were positively responded to
microorganisms inoculation than other treatments.
Leaves of both cultivars showed significant increase of N, K, Ca and Mg when grown
nd
on peatmoss in 2 season. Neither cultivar nor substrate as single factors affected vitamin C
or TSS content, with some exceptions. Dry matter, nitrate and total sugar in fruits of cultivars
nd
did not differ under both substrates, with some exceptions in 2 season. However, inoculation
of plants with Glomus + Azospirillum improved vitamin C contents in fruits of both cultivars,
especially when cucumbers grown on peatmoss. Similarly, TSS content was significantly
higher in fruits of cucumbers grown in peatmoss, but when inoculated with Glomus +
Azotobacter + Azospirillum. In this respect, it could be said that microorganisms inoculation
improved nutrients content in cucumber fruits if compared to the control. This conclusion
based on fact that microorganisms play a vital role to increase nutrients contents and plant
metabolism processes in leaves and shoots of inoculated plants.
V. ZÁV ĚR
Práce probíhala v letech 2006 a 2007 v otevřeném hydroponickém systému ve skleníku
Zahradnické fakulty v Lednici, Mendelovy Univerzity v Brně, Česká Republika. Rostliny
okurek (Cucumis sativus L. odrůdy ‘Passandra’ a ‘Girola’) byly vysázeny ve 4 opakováních ve
split-plot designu. Rostliny byly pěstovány včedičové plsti a v rašelině.
Sadba byla inokulována ětmito mikroorganismy: N1 (Azotobacter chroococcum), N2
(Azospirillum brasilense), G (Glomus mosseae), a jejich kombinace N1 + N2, G + N1, G +
N2, G + N1 + N2 a kontrola.
Obecně zhodnoceno, odrůda Passandra vykázala vyšší výšku rostliny, po čet listů,
obsah čerstvé a suché hmoty v listech. Na druhé straněodrůda Girola dosáhla ve 2. roce
průkazně vyšší rozsah listové plochy. Ranost a celkový výno s byl u obou odrůd srovnatelný.
Tyto výsledky ukazují vliv genotypu na r ůstové parametry. Přitom Girola dosáhla průkazně
vyšší délky plodů, jejich průměru a hmotnosti. Passandra ovšem vykazovala v obou s ezónách
průkazně vyšší po čet plodů na rostlinu a současně vyšší zastoupení jakosti „Výb ěr“ v roce
2007.
Dusík, fosfor a hořčík nevykazoval rozdílný obsah v listech obou odr ůd. Obsah
draslíku byl průkazně vyšší u odr ůdy Girola v roce 2006 a naopak u odrůdy Passandra v roce
2007. Obsah dusíku v plodech odrůdy Passandra a P, K, Ca v plodech odrůdy Girola byl vyšší
v roce 2006. Genotyp neovlivnil vitamin C ani TSS obsah. Obsah sušiny v plodech odr ůdy
Passandra byl průkazně vyšší v obou letech; celkový obsah cukr ů v roce 2006. Obsah
dusičnanů byl průkazně vyšší v plodech odr ůdy Girola, ale stále pod hygienickým limitem.
Lze shrnout, že rozdíly mezi odr ůdami korelovaly s rozdíly danými jejich genotypem. Odrůdy
se lišily v produktivit ě výnosu a kvalit ě.
Vliv rozdílných substrátů přinesl poznatek, že rostliny p ěstované v rašelinovém
substrátu dosáhly ůprkazně vyššího po čtu listů na rostlinu a obsahu sušiny v listech. Naopak
listová plocha ičerstvá hmotnost listů byla mezi sezónami variabilní. Mírný pr ůkazný nárůst
byl zjišt ěn v případě ranosti rostlin pěstovaných v čedičové plsti. Oproti tomu byl zjištěn
průkazně vyšší výnos – v roce 2006 v rašelinovém a v roce 2 007 v čedičovém substrátu.
Rostliny v čedičové plsti dosáhly průkazně vyšší délky a hmotnosti. Ovšem okurky z
rašelinového substrátu dosahovaly průkazně vyšší po čet plodů na rostlinu a zastoupení „Výb
ěru“ v roce 2006, na rozdíl od výsledku v další sezó ně. Obsah minerálních látek, vitaminu C,
cukrů i TSS v plodech nebyl mezi substráty rozdílný, s několika výjimkami. Plody rostlin z
-
čedičové plsti dosáhly průkazně vyšší obsah sušiny a NO 3 v roce 2006.
Z tohoto hlediska lze shrnout, že rozdíl v látkovémsložení plod ů závisí jen zčásti na použitém
pěstebním substrátu.
Rostliny inokulované N-Fixujícími bakteriemi nebo houbou Glomus nevykazovaly
rozdíly v růstu, ranosti, délce a průměru plodů, počtu plodů na rostlinu v obou letech, s
několika výjimkami, oproti neošet řené kontrole. Na druhé straně inokulace pozitivně přispěla
k zlepšení r ůstových parametr ů a výnosu, i když výsledky nebyly signifikantní. Je
třeba poukázat na fakt, že mikroorganismy byly vystaveny v hydroponickém systémuřadě
nepříznivých faktor ů, jako kyselé pH, dostatečný obsah živin v živném roztoku, což mohlo
vést ke snížení efektu inokulace.
Dále také, všechna inokulovaná ošetření vedla k vyššímu, i když nepr ůkaznému zvýšení
výnosu. Toto navýšení dosáhlo 8 a 21 % v 1., respe ktive 2. sezón ě. Také bylo doloženo
signifikantní zvýšení pr ůměrné hmotnosti plodů rostlin inokulovaných samostatným
Azotobacter nebo v kombinaci s Azospirillium, ve srovnání s kontrolou v roce 2006. Inokulace
rostlin vedla k zvýšení tržního výnosu, o bsahu makroprvků v listech a plodech obou odrůd, s
několika výjimkami.
Obsah vitaminu C v plodech byl průkazně nejvyšší p ři inokulaci Glomus + Azospirillum.
Také obsah celkových rozpustných solí v plodech byl vyšší p ři inokulaci Glomus +
Azotobacter + Azospirillum v roce 2006. Na druhé straně průkazné snížení sušiny plod ů bylo
zjišt ěno ve variantě Azotobacter a Azospirillum v roce 2007, ve srovnání s kontrolou. Obsah
nitrátů byl průkazně zvýšen p ři inokulaci Azotobacter nebo Azospirillum oproti variantě
Glomus nebo kontrole v roce 2006.
Interakce mezi odrůdou, substrátem a inokulací ukázalyřadu kladných vliv ů na většinu
růstových charakteristik rostlin okurek. Kup říkladu N-Fixers samostatně nebo v kombinaci s
Glomus zvýšily r ůst rostlin v rašelinovém substrátu. Okurky v rašelině inokulovanéGlomus +
Azotobacter nebo Azospirillum měly průkazně vyšší celkový výnos v roce 2006 oproti
kontrole. Lze shrnout, že výnos a r ůst odrůdy Passandra v rašelinovém substrátu byl pozitivně
ovlivněn inokulací mikroorganismy.
Listy u obou odrůd dosáhly průkazně vyššího obsahu N, K, Ca a Mg p ři pěstování v
rašelin ě v roce 2007. Ani odrůda ani substrát jako samostatný faktor neměly vliv na obsah
vitaminu C nebo TSS, mimo několika výjimek. Obsah sušiny, nitrátů a celkových cukr ů v
plodech odrůd nebyl rozdílný mezi ob ěma substráty, mimo několika případů v roce 2007.
Ovšem inokulace s Glomus + Azospirillum zvýšila obsah vitaminu C v plodech obou odr ůd,
především v rašelinovém substrátu. Podobně, TSS obsah byl průkazně vyšší v plodech rostlin z
rašeliny, pokud byly rostliny inokulovány Glomus + Azotobacter + Azospirillum. Z tohoto
hlediska lze konstatovat, že inokulace zlepšila nut riční hodnotu plodů oproti neošet řené
kontrole. Toto konstatování odpovídá faktu, že symbiotické mikroorganismy mají podpůrnou
roli v rámci rostlinného metabolismu v listech a výhonech inokulovaných rostlin.
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VII. APPENDIX
Table 9. Effect of interaction between cultivar x microorganisms inoculation on plant
height (m) of cucumbers in 2006 and 2007
Table 10. Effect of interaction between substrate x microorganisms inoculation on plant
height (m) of cucumbers in 2006 and 2007
Table 11. Effect of interaction between cultivar x microorganisms inoculation on leaf
2
area (cm ) of cucumbers in 2006 and 2007
Table 12. Effect of interaction between substrate x microorganisms inoculation on leaf
2
area (cm ) of cucumbers in 2006 and 2007
Table 13. Effect of interaction between cultivar x microorganisms inoculation on number
of leaves of cucumbers in 2006 and 2007
Table 14. Effect of interaction between substrate x microorganisms inoculation on
number of leaves of cucumbers in 2006 and 2007
Table 15. Effect of interaction between cultivar x microorganisms inoculation on fresh
weight (g/leaf) of cucumbers in 2006 and 2007
Table 16. Effect of interaction between substrate x microorganisms inoculation on fresh
weight (g/leaf) of cucumbers in 2006 and 2007
Table 17. Effect of interaction between cultivar x microorganisms inoculation on dry
weight (g/leaf) of cucumbers in 2006 and 2007
Table 18. Effect of interaction between substrate x microorganisms inoculation on dry
weight (g/leaf) of cucumbers in 2006 and 2007
Table 21. Effect of interaction between cultivar x microorganisms inoculation on early
-2
yield (kg.m ) of cucumbers in 2006 and 2007
Table 22. Effect of interaction between substrate x microorganisms inoculation on early
-2
yield (kg.m ) of cucumbers in 2006 and 2007
Table 23. Effect of interaction between cultivar x microorganisms inoculation on total
-2
yield (kg.m ) of cucumbers in 2006 and 2007
Table 24. Effect of interaction between substrate x microorganisms inoculation on total
-2
yield (kg.m ) of cucumbers in 2006 and 2007
Table 27. Effect of interactions between cultivar x microorganisms inoculation on fruit
length (mm) of cucumbers in 2006 and 2007
Table 28. Effect of interactions between substrate x microorganisms inoculation on fruit
length (mm) of cucumbers in 2006 and 2007
Table 29. Effect of interactions between cultivar x microorganisms inoculation on fruit
diameter (mm) of cucumbers in 2006 and 2007
Table 30. Effect of interactions between substrate x microorganisms inoculation on fruit
diameter (mm) of cucumbers in 2006 and 2007
Table 31. Effect of interactions between cultivar x microorganisms inoculation on fruit
number per plant of cucumbers in 2006 and 2007
Table 32. Effect of interactions between substrate x microorganisms inoculation on fruit
number per plant of cucumbers in 2006 and 2007
Table 33. Effect of interactions between cultivar x microorganisms inoculation on fruit
weight (g) of cucumbers in 2006 and 2007
Table 34. Effect of interactions between substrate x microorganisms inoculation on fruit
weight (g) of cucumbers in 2006 and 2007
Table 37. Effect of interaction between cultivar x microorganisms inoculation on fruit
-2
yield of grade "Extra" (kg.m ) in 2006 and 2007
Table 38. Effect of interaction between substrate x microorganisms inoculation on fruit
-2
yield of grade "Extra" (kg.m ) in 2006 and 2007
Table 39. Effect of interaction between cultivar x microorganisms inoculation on fruit
-2
yield of "Class A" (kg.m ) in 2006 and 2007
Table 40. Effect of interaction between substrate x microorganisms inoculation on fruit
-2
yield of "Class A" (kg.m ) 2006 and 2007
Table 41. Effect of interaction between cultivar x microorganisms inoculation on fruit
-2
yield of "Class B" (kg.m ) in 2006 and 2007
Table 42. Effect of interaction between substrate x microorganisms inoculation on fruit
-2
yield of "Class B" (kg.m ) in 2006 and 2007
Table 43. Effect of interaction between cultivar x microorganisms inoculation on
-2
unmarketable fruit yield (kg.m ) in 2006 and 2007
Table 44. Effect of interaction between substrate x microorganisms inoculation on
-2
unmarketable fruit yield (kg.m ) in 2006 and 2007
Table 47. Effect of interaction between cultivar x microorganisms inoculation on nitrogen
content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 48. Effect of interaction between substrate x microorganisms inoculation on
nitrogen content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 49. Effect of interaction between cultivar x microorganisms inoculation on
phosphorus content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 50. Effect of interaction between substrate x microorganisms inoculation on
phosphorus content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 51. Effect of interaction between cultivar x microorganisms inoculation on
-1
potassium content (mg.kg fresh weight) of cucumbers leaves in 2006 and
2007
Table 52. Effect of interaction between substrate x microorganisms inoculation on
-1
potassium content (mg.kg fresh weight) of cucumbers leaves in 2006 and
2007
Table 53. Effect of interaction between cultivar x microorganisms inoculation on calcium
-1
content (mg.kg fresh weight) of cucumbers leaves in 2006 and 2007
Table 54. Effect of interaction between substrate x microorganisms inoculation on
-1
calcium content (mg.kg fresh weight) of cucumbers leaves in 2006 and 2007
Table 55. Effect of interaction between cultivar x microorganisms inoculation on
-1
magnesium content (mg.kg fresh weight) of cucumbers leaves in 2006 and
2007¨
Table 56. Effect of interaction between substrate x microorganisms inoculation on
-1
magnesium content (mg.kg fresh weight) of cucumbers leaves in 2006 and
2007
Table 59. Effect of interaction between cultivar x microorganisms inoculation on nitrogen
content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 60. Effect of interaction between substrate x microorganisms inoculation on
nitrogen content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 61. Effect of interaction between cultivar x microorganisms inoculation on
phosphor content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 62. Effect of interaction between substrate x microorganisms inoculation on
phosphor content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 63. Effect of interaction between cultivar x microorganisms inoculation on
-1
potassium content (mg.kg fresh weight) of cucumber fruits in 2006 and 2007
Table 64. Effect of interaction between substrate x microorganisms inoculation on
potassium content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and 2007
Table 65. Effect of interaction between cultivar x microorganisms inoculation on calcium
-1
content (mg.kg fresh weight) of cucumber fruits in 2006 and 2007
Table 66. Effect of interaction between substrate x microorganisms inoculation on
-1
calcium content (mg.kg fresh weight) of cucumber fruits in 2006 and 2007
Table 67. Effect of interaction between cultivar x microorganisms inoculation on
-1
magnesium content (mg.kg fresh weight) of cucumber fruits in 2006 and 2007
Table 68. Effect of interaction between substrate x microorganisms inoculation on
-1
magnesium content (mg.kg fresh weight) of cucumber fruits in 2006 and 2007
Table 71. Effect of interaction between cultivar x microorganisms inoculation on vitamin
-1
C (mg.kg ) of cucumber fruits in 2006 and 2007
Table 72. Effect of interaction between substrate x microorganisms inoculation on
-1
vitamin C (mg.kg ) of cucumber fruits in 2006 and 2007
Table 73. Effect of interaction between cultivar x microorganisms inoculation on TSS
(%) of cucumber fruits in 2006 and 2007
Table 74. Effect of interaction between substrate x microorganisms inoculation on TSS
(%) of cucumber fruits in 2006 and 2007
Table 75. Effect of interaction between cultivar x microorganisms inoculation on dry
matter (%) of cucumber fruits in 2006 and 2007
Table 76. Effect of interaction between substrate x microorganisms inoculation on dry
matter (%) of cucumber fruits in 2006 and 2007
Table 77. Effect of interaction between cultivar x microorganisms inoculation on NO3-
-1
(mg.kg ) of cucumber fruits in 2006 and 2007
Table 78. Effect of interaction between substrate x microorganisms inoculation on NO3-
-1
(mg.kg ) of cucumber fruits in 2006 and 2007
Table 79. Effect of interaction between cultivar x microorganisms inoculation on total
-1
sugars (mg.kg ) of cucumber fruits in 2006 and 2007
Table 80. Effect of interaction between substrate x microorganisms inoculation on total
-1
sugars (mg.kg ) of cucumber fruits in 2006 and 2007