You are on page 1of 54

IMPACT OF AVOCADO (Persea americana Mill.

) TREE CANOPY
MANAGEMENT AND HONEY BEE POPULATION DENSITY ON
AVOCADO FRUIT AND HONEY PRODUCTION
BY:

TESEMA YOHANIS

SUPERVISORS
1. Dr. Meseret Tesema (PhD)
2. Dr. Alemu Dessa (PhD)

A THESIS PROPOSAL SUBMITTED TO THE SCHOOL OF GRADUATE


STUDIES OF HAWASSA UNIVERSITY

IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE


OF MASTER OF SCIENCE IN PLANT AND HORTICULTURAL SCIENCE
(SPECIALIZATION: HORTICULTURE)

AUGUST, 2021
HAWASSA, ETHIOPIA
APPROVAL SHEET

HAWASSA UNIVERSITY COLLEGE OF AGRICULTURE

SCHOOL OF PLANT AND HORTICULTURAL SCIENCE

Thesis proposal Approval sheet

The Research Proposal Entitled as “Impact of Avocado Tree Canopy Management and Honey
Bee Population Density on Avocado Fruit and Honey Production
In case of Tereri Kela, Sidama Regional State, Ethiopia”. Has been approved by school of plant
and horticultural science for partial fulfillment of The Requirements for the Degree of Master of
Science in plant and horticultural science (Specialization: Horticulture)

Submitted by:-

Tesema Yohanis
------------------------------------------------- ------------------- -------------------------
Name of student Signature Date

Approved by:-
Dr. Meseret Tesema(PhD)
---------------------------------------------------- -------------- -----------------------------
Name of major advisor Signature Date

Dr. Alemu Dessa (PhD)


---------------------------------------------------- ---------------- -----------------------------
Name of co-advisor Signature Date

---------------------------------------------------- ------------------ ----------------------------


Name of school head Signature Date

----------------------------------------------------- -------------------- ---------------------------


School of graduate studies Signature Date

ii
Tables..............................................................................................................................................iv

1. INTRODUCTION.......................................................................................................................1

1.1. Background and Justification................................................................................................1

1.2. Statement of Problem............................................................................................................4

1.3. Objective of the Study...........................................................................................................5

1.4. Research Question.................................................................................................................5

2. LITERATURE REVIEW............................................................................................................6

2.1. The avocado fruit tree...........................................................................................................6

2.1.1. History and Distribution.................................................................................................6

2.1.2. Ecology of avocado fruit tree: Climate and Soils...........................................................6

2.1.3. Global Avocado Production...........................................................................................8

2.1.3.1. World avocado production......................................................................................8

2.1.3.2. Avocado production in Africa.................................................................................9

2.1.3.3. Avocado production in Ethiopia............................................................................10

2.1.4. Avocado fruit Uses and Composition..........................................................................11

2.2. Avocado fruit tree and canopy management.......................................................................13

2.3 Avocado fruit tree light interception and light use efficiency.............................................14

2.3.1 Light interception and photosynthesis of tree canopy...................................................14

2.3.2. The Avocado fruit trees response to light....................................................................15

2.4 Avocado tree flower, pollination mode and pollinizers..................................................16

2.4.1. Avocado tree flower and Flowering.............................................................................16

2.4.1.1. Avocado tree flower..............................................................................................16

2.4.1.2 Avocado tree flowering..........................................................................................17

2.4.2. Avocado tree pollination Modes..................................................................................18

iii
2.4.2.1. Cross-pollination....................................................................................................18

2.4.2.2. Close-pollination....................................................................................................18

2.4.2.3. Self-pollination......................................................................................................19

2.4.3. Avocado tree pollinizer................................................................................................19

2.5. Avocado fruit orchard and Honey bee population..............................................................20

3. MATERIAL AND METHODS.................................................................................................22

3.1. Description of the Study Area.............................................................................................22

3.2. Experimental Design and procedure...................................................................................22

3.3. Data collection....................................................................................................................23

3.4 Statistical Analysis.............................................................................................................24

4. WORK PLAN............................................................................................................................25

5. BUDGET BREAKDOWN........................................................................................................26

6. REFERENCE............................................................................................................................28

iv
List of Tables
Table 2.1. Avocado production of the top 12 producing countries in 2000 and 2008....................9
Table 2.2. Africa top 10 avocado producing countries (2008-2012, in mt)....................................9
Table 4.1. Work plan schedule......................................................................................................25
Table 5.1. Stationery.....................................................................................................................26
Table 5.2. Transport Cost.............................................................................................................26
Table 5.3. Farm and laboratory inputs...........................................................................................26
Table 5.4. Labor cost....................................................................................................................27
Table 5.5. Personnel cost...............................................................................................................27

v
1. INTRODUCTION

1.1. Background and Justification


The avocado (Persea americana Mill.) is an evergreen subtropical fruit tree native to Central
America and Mexico (Chen et al., 2009). It contains many vitamins, fat-soluble, protein,
potassium and unsaturated fatty acid that are less common in other fruits and it is used in the
pharmaceutical and cosmetic industries as a raw material. (Duarte et al., 2017). Total world
production is more than 4.5 million tones, with about 25% of the crop traded around the globe
(Bost et al., 2013). Mexico is the largest producer, with a total production of about 1.5 million
tones (28% of world production). Other important producing countries are Chile (8%),
Dominican Republic (7%), Indonesia (6%), Colombia (5%), Peru (5%), United States (5%),
Brazil (4%), Kenya (4%) and Rwanda (3%).
Avocado is a subtropical to tropical tree, and can be grown successfully from the tropics to the
subtropics at latitude of 35°. There is a wide variation in the performances of the three races and
various cultivars in different growing environments. Avocado trees are variable in shape, from
tall upright specimens to widely spreading forms with multiple branches (Chanderbali et al.,
2013). The avocado trees can grow to heights of 15–18 m, with commercial orchards usually
pruned to some degree.
Avocado was first introduced to Ethiopia in 1938 by private orchardists in Hirna and Wondo-
genet (Edossa, 1997; Woyessa and Berhanu, 2010; and Zekarias, 2010). Annual avocado
production in Ethiopia is 25633.16 tons. The production and productivity of avocado in Ethiopia
remained low compare to world average.
The low production and productivity of avocado in Ethiopia in general and in study area are the
results of various factors. Among which lack of improved varieties, poor canopy management
practices, pests and diseases and low pollinator visits or lack of efficient pollinators are
important factors (MoA, 2005).

In
recognition of its nutritional quality
and economic
1
importance, the crop has been
distributed to the rest of
the world since the last 400 years.
During the last
century, it has been widely distributed
to more than 50
countries around the world, which
mainly includes
sub-tropical and tropical areas such as
sub-Saharan
Africa. Ethiopia is one of the top five
avocado
producers in sub-Saharan Africa.
Avocado was first
introduced to Ethiopia around 1938 by
private
orchardists in Hirna (Eastern
highlands of Ethiopia)
2
and Wondo-genet (Southern
Highlands of Ethiopia).
Gradually, it has been distributed to
different agro-
ecologic conditions where the crop
could be adapted
(Edossa 1997; Megersa and Alemu
2013; Woyessa
and Berhanu 2010). Despite its long
history since
introduction and the diverse agro-
ecologic conditions
of Ethiopia, its distribution is still
limited to few areas
of the country (CSA 2014). Currently
the main
avocado producing areas include
Sidama and Wolay-
3
ita areas in the South, Jimma and
Mizan areas in south-
western and Hararge area in eastern
region of the
country (CSA 2014; Megersa and
Alemu 2013).
In Ethiopia, avocado trees are mainly
grown as an
integral component of coffee (Coffea
Arabica L.) and
enset (Ensetventricosum) agroforesrty
systems and to
a lesser extent in combination with
other crops like
maize (Moges 2009; Shumeta 2010).
As coffee
requires shade trees and known for its
shade-tolerance,
4
smallholder producers consider
avocado as one of the
important shade trees while also
benefiting a lot from
the sale of the avocado fruits
(Kahuranga et al. 1993;
Moges 2009; Shumeta 2010). It is also
grown as
individual trees around the court yards
where it is used
also as shades for people and
livestock.
The Sidama Highland in Southern
Ethiopia is known
for the traditional coffee- and enset-
based agroforestry
system (Abebe et al. 2009; Asfaw and
Agren 2007;
5
Asfaw and Lemenih
2010; M o g e s 2009). This zone is
one of the areas where avocado was
first introduced to
Ethiopia (Megersa and Alemu
2013; Z e k a r i a s 2010).
The Sidama highlands have suitable
agro-ecological
conditions for rainfed avocado
production. It accounts
about 36 % of the national annual
avocado production
(CSA 2014). Therefore, the Sidama
highlands are
important sources of avocado fruits for
major cities
such as Hawassa and Addis Ababa
In
6
recognition of its nutritional quality
and economic
importance, the crop has been
distributed to the rest of
the world since the last 400 years.
During the last
century, it has been widely distributed
to more than 50
countries around the world, which
mainly includes
sub-tropical and tropical areas such as
sub-Saharan
Africa. Ethiopia is one of the top five
avocado
producers in sub-Saharan Africa.
Avocado was first
introduced to Ethiopia around 1938 by
private
7
orchardists in Hirna (Eastern
highlands of Ethiopia)
and Wondo-genet (Southern
Highlands of Ethiopia).
Gradually, it has been distributed to
different agro-
ecologic conditions where the crop
could be adapted
(Edossa 1997; Megersa and Alemu
2013; Woyessa
and Berhanu 2010). Despite its long
history since
introduction and the diverse agro-
ecologic conditions
of Ethiopia, its distribution is still
limited to few areas
of the country (CSA 2014). Currently
the main
8
avocado producing areas include
Sidama and Wolay-
ita areas in the South, Jimma and
Mizan areas in south-
western and Hararge area in eastern
region of the
country (CSA 2014; Megersa and
Alemu 2013).
In Ethiopia, avocado trees are mainly
grown as an
integral component of coffee (Coffea
Arabica L.) and
enset (Ensetventricosum) agroforesrty
systems and to
a lesser extent in combination with
other crops like
maize (Moges 2009; Shumeta 2010).
As coffee
9
requires shade trees and known for its
shade-tolerance,
smallholder producers consider
avocado as one of the
important shade trees while also
benefiting a lot from
the sale of the avocado fruits
(Kahuranga et al. 1993;
Moges 2009; Shumeta 2010). It is also
grown as
individual trees around the court yards
where it is used
also as shades for people and
livestock.
The Sidama Highland in Southern
Ethiopia is known
for the traditional coffee- and enset-
based agroforestry
10
system (Abebe et al. 2009; Asfaw and
Agren 2007;
Asfaw and Lemenih
2010; M o g e s 2009). This zone is
one of the areas where avocado was
first introduced to
Ethiopia (Megersa and Alemu
2013; Z e k a r i a s 2010).
The Sidama highlands have suitable
agro-ecological
conditions for rainfed avocado
production. It accounts
about 36 % of the national annual
avocado production
(CSA 2014). Therefore, the Sidama
highlands are
important sources of avocado fruits for
major cities
11
such as Hawassa and Addis Ababa
Due to its flowering and fruiting characteristics the avocado tree must produce new growth each
year to remain productive (Whiley and Schaffer, 1994). However, if left unchecked the orchard
eventually becomes crowded with a loss in yield and fruit quality. The basic problem with
overcrowded orchards is insufficient light penetration through the canopy (Stadler and Stassen,
1985). There are several systems to manage tree size and improve light interception and
penetration, including tree thinning, selective limb removal and mechanical pruning. Significant
advances have been made in the development of mechanized pruning and growth retardants in
avocado orchards. However, recommendations on canopy management strategies for Ethiopian
growers have not yet been identified.

Sunlight is a resource that is largely taken for granted in most agricultural systems. Fruit
production involves the capture and conversion of radiant/light energy into chemical energy in
fruit biomass (dry matter). The main controlling factors here are the amount of incoming
radiation, the percentage of that radiation which is intercepted by the tree leaves, and how
efficiently the tree converts that energy into fruit (Wünche and Lakso, 2000). Through the
process of photosynthesis, light energy is converted into stored energy (biomass) in the form of
vegetative and reproductive structures. The efficient capture and conversion of light is integral
to high yields. Potential light interception firstly is determined by the amount of light
interception and conversion efficiency. The more light there is, the higher the proportional
amount that can be intercepted. Distributed light is the light that is transmitted through the
canopy and as such is a function of tree and orchard factors. Of these factors, tree architecture,
leaf area index, row spacing and row direction/orientation are the most important (Palmer, 2004).
Monteith (1977) demonstrated a fundamental correlation between crop dry matter production
and seasonal accumulated light interception. While there have been few avocado studies on this
matter, a number of studies in apple orchards (Lakso, 1994) have shown that there is a close
positive relationship between total seasonal light interception and fruit yield. In a four-year study
of apple orchards in Australia, Middleton (2007) concluded that the most productive orchards
had LAIs of between two and three. Productivity declined as LAI fell below 1.5 due to the trees
not having enough canopy volume for high light interception. Further, Middleton (2007) showed
that yields increased as light interception increased from 55 to 62% but the majority of orchards

12
surveyed intercepted less than this due to inadequate Tree Row Volume as a result of being
planted too far apart within rows and inter-rows being wider than necessary (i.e. planting density
was not high enough). In the same study, tree height as a parameter of architecture, was found to
be of less importance than LAI. In many orchard crops, light penetration is important for the
development of quality characteristics in apples (Palmer, 2004) and it is also linked with flower
bud development (Palmer, 2004), dormancy, shoot growth and leaf morphology (Bastias &
Corelli-Grappadelli, 2012) in addition to its role in assimilation. Shading within the canopy has
direct detrimental effects on assimilation when shaded leaves become sinks for energy due to a
negative net photosynthetic rate and are effectively parasitic (Hadari, 2004). In avocado, Hadari
(2004) showed that maximum light penetration through a healthy canopy was 1m, suggesting
that canopy management should aim at restricting the thickness of the foliage.
Avocado is one of at least 105 crops that receive yield benefits from animal pollination (Rader et
al., 2020), and together, these crops represent approximately 35% of total agricultural production
(Klein et al., 2007).Inadequate pollination has been suggested as an important factor in the
limiting of the avocado (Persea americana Mill.) yield. This species presents the phenomenon of
dichogamy proterogyny synchrony, i.e. sequential development of reproductive functions.
Avocado cultivars are classified into complementary flowering groups A and B, based on their
daily flowering pattern. The type A flowers open in the female form in the morning and close in
the afternoon but then open again in the masculine form in the afternoon of the following day.
The type B flowers open in the female form in the afternoon; they then close and reopen the
following morning in the male form (Sedgley, 1979; Ish-Am, 2004). The adoption of dichogamy
as a breeding strategy implies that for an effective transfer of pollen, insects should visit the
flowers in both sexual states. Fruit set is minimal or absent when insect pollinators such as bees
and flies are excluded through caging (Malerbo-Souza et al., 2000). Pollen-carrying vectors play
a key role in the pollination and increase in the genetic variability of avocados because a
considerable number of fruits are produced only through cross pollination (Bergh, 1977; Gazit &
Degani, 2002). Avocado flowers are visited by a variety of insects including bees, flies, wasps,
beetles and thrips (Vithanage, 1990). The honey bee A. mellifera is considered the main
pollinating agent of the avocado (Free, 1970; Nieto, 1984; Davenport, 1986; Ish-Am &
Eisikowitch, 1993; Avilán & Rodríguez, 1995; Castañeda et al., 1999; Peña, 2003; Can-Alonso
et al., 2005; Goodwin, 2012). However, despite this and in comparison with other fruit trees,

13
bees do not work efficiently in avocado flowers as they get more attracted by flowers of wild
plants grown in orchards (Ish-Am & Eisikowitch, 1993). An avocado tree produces about one
million flowers and 10000 to 40000 female flowers open each day in Israel (Lahav & Zamet,
1999). A good seasonal crop of 400 to 600 fruits per tree requires the pollination and fertilization
of about the same number of flowers, which may be accomplished with only two or three forager
honey bees. However, in practice a measurable initial fruit set under field condition demands the
work of at least five to ten honey bees per tree throughout the female bloom (Ish-Am, 2004). In
avocado orchards, it is common practice to introduce colonies of bees to promote pollination
(Pérez-Balam et al., 2012). In New Zealand, four to ten hives/ha is recommended (Evans,
Goodwin, & Mcbrydie, 2010), while in Israel, it the optimal number of hives/ha has been found
to be eight, placed at distances not less than 100 m between them (Bergh, 1977). Goodwin
(2012) recommended the introduction of hives when the crop presented between 5 and 10 % of
flowering, guaranteeing its persistence in the flowers. Honey bee pollinator number density is the
most important of the several factors that contribute and change to increase productivity.
Research has shown that pollination by honey bee increases the fruit set in avocados (Ish-Am &
Eisikowitch, 1993; Goodwin, 2012).

1.2. Statement of Problem


Horticulture can be an important factor for economic development and contribute to increased
food security and improve the populations’ nutrition intake (Weinberger & Lumpkin, 2007). The
growing population and changing dietary habits in Ethiopia has increased the demand for fruit
(ILRI, 2011). Avocado is one of the most important fruit crops cultivated mostly in Tefer kella
district in Sidama Region. However, its productivity is remained very low compared to the world
average. The reason of low productivity of avocado is associated with poor canopy management
and low Pollinator visits and lack of efficient pollinators, among other factors (MoA, 2005). Poor
canopy manipulation is the major issues to reducing the avocado fruiting. Due to its flowering
and fruiting characteristics the avocado tree must produce new growth each year to remain
productive (Whiley and Schaffer, 1994). However, if left unchecked the orchard eventually
becomes crowded with a loss in yield and fruit quality. The basic problem with overcrowded
orchards is insufficient light penetration through the canopy (Stadler and Stassen, 1985).
Avocado is a tree that mainly relies on insect pollinators for formation of fruits and viable seeds
(Peña et al., 2002; Ish-Am, 2004). Inadequate pollination has been suggested as an important
14
factor in the limiting of the avocado (Persea americana Mill.) yield. The honey bee A. mellifera
is considered the main pollinating agent of the avocado (Peña, 2003; Can-Alonso et al., 2005;
Goodwin, 2012). Poor canopy manipulation and insufficient pollination are the major issues to
reducing the avocado fruiting. This issue has been raising concerns that the livelihoods of
avocado farmers will be badly affected. Despite these concerns, there is scarcity of information
on avocado canopy manipulation for light interception in the avocado fruit tree and hive bee
population density on pollination of avocado fruit tree in study area and in Ethiopia are not well
understood. Therefore, this study deals with avocado canopy management to attain good sunlight
distribution through the avocado tree canopy for better fruiting and honey bee visitation activity
per tree will be evaluates and recommends avocado manipulation method and optimum honey
bee density per tree for better avocado tree productivity at study area.

1.3. Objective of the Study


General Objective
 To evaluate avocado canopy management, manipulation sunlight distribution and honey
bee population on avocado fruiting at Sidama Region Tefer Kela District.

Specific Objectives
 To evaluate the effect of canopy management on sun light interception and the avocado
trees yield
 To determine the appropriate canopy size/management methods for efficient light
distribution and avocado tree productive
 To determine contribution of honey bee population on efficient pollination, fertilization
and fruit set of the avocado
 To determine the honey bee hive population/density per avocado tree number for
effective fruit and honey production

1.4. Research Question


a) What is the effect of canopy management to maximize avocado fruiting at study area?
b) How can avocado canopy manages to attain good sunlight interception through avocado
fruit trees for better fruiting?

15
c) What is the effect of honey bee population on avocado pollination and fruiting at study
area?
d) How can honey bee pollination be improved for higher avocado fruiting?
e) What are the current evidence gaps and what should be the focus of future research?

2. LITERATURE REVIEW

2.1. The avocado fruit tree

2.1.1. History and Distribution


The avocado (Persea americana Mill.) is a polymorphic tree species that originated in a broad
geographical region stretching from the Pacific coast of Central America through Guatemala to
the eastern and central highlands of Mexico (Popenoe, 1920). Three distinct and separate taxa or
sub-species now termed the Guatemalan, Mexican and West Indian or Antillean races have been
selected over millennia (Knight, 2002). Archaeological evidence shows that when the climatic
conditions changed during the Paleocene glaciations, avocado ancestors migrated from North
America to the south and became established in the more hospitable habitats of Mesoamerica
(Schroeder, 1968; Storey et al., 1986; Scora and Bergh, 1992; Bergh, 1995). Evidence suggests
that the complex geological history of Mexico has been the main evolutive factor for the avocado
(Ramamoorthy et al., 1993). The avocado has been consumed in Mesoamerica by human groups
since prehistoric times (Mac Neish, 1964). Buckler et al. (1998) documented that from 16,000 to
8,000 BC the weather in this region was appropriate for avocado development. Today the
avocado is cultivated all over the world. In 1856 avocados were brought to California by
Nicaraguan settlers. In Israel, the first avocado was introduced in 1908. From 1933 to 1998
avocado selections were introduced in Central America, the Caribbean, North and South
America, Africa, Asia, Oceania, and Europe (Knight, 2002). In recent years, the avocado has
become the fourth most important tropical fruit in the world, and it continues to increase in
importance in many places (Bergh, 1992). China, for example, produced 45,000 tons in 1996,
whereas in 1991 there was no avocado production reported (Knight, 2002). At present, Mexico is
the main producer (Foreign Agricultural Service, USDA, 2006) with 1,021,515 tons in 2005
(SIAP, 2007). Current varieties and rootstocks for avocado cultivation in the world are the

16
products of various breeding programs based on exploration, collections, conservation, and
evaluation trials throughout their regions of origin and dispersion (Mijares and López, 1998).

2.1.2. Ecology of avocado fruit tree: Climate and Soils


Avocado first became an important orchard crop early in the twenty century. It is therefore a
relatively new or ‘young’ crop, at an early stage of domestication. It possesses residual eco
physiological features that are adaptive to competing in a neotropical rainforest (often at high
altitude), which can be counter-productive to the needs of modern orcharding (Wolstenholme,
1986; Wolstenholme and Whiley, 1998, 1999). Mexican ecotypes are reported to be indigenous
to highland areas between 19° and 24°N, i.e. borderline subtropical highland to semi-tropical
highland (Storey et al., 1986). Wild Guatemalan avocados range from 14–16°N in mountainous
terrain, i.e. truly tropical highland. The most tropical ecotype, the West Indian or Lowland
(Antillean), extends from 8–15°N in lowland coastal areas on the Pacific coast, and the term
West Indian is a misnomer. A latitudinal range of 8–24°N and altitudes from sea level to over
2500 m is therefore presumed for wild avocados. In contrast, avocados are cultivated
commercially from about 40°N on the coast of the Black Sea in the Batoum region (Gaillard,
1987), to the Bay of Plenty on the North Island in New Zealand (c.39°S), a huge latitudinal
spread. Praloran (1970) summarized climatic data for the three ‘races’ of avocado. Three
representative climate types were chosen: the Mexican highlands at 1400–2700 m; the
Guatemalan highlands at 1500–2350 m; and the Colombian lowlands at 100–450 m (today not
considered as the centre of origin of the West Indian race, but representative of a tropical
lowland climate). Praloran (1970) concluded that a common feature of indigenous avocado areas
was the marked dry season in which flowering took place. Abundant rain during flowering (as in
the humid tropics) results in a prolongation of the flowering period. Flowering often lasts 4–6
months in the humid tropical lowlands of the Ivory Coast and Cameroon, which indicates weak
climatic synchronization of flowering and may be a response to high temperatures. Mexican and
Guatemalan ecotypes therefore have a more synchronized flowering period (about 1 month)
when flowering occurs in a cool (mean 12.7–21°C) and dry period. West Indian ecotypes are
better adapted to hot and relatively dry (but with high relative humidity) conditions during
flowering. Praloran (1970), citing the pollination study of Lesley and Bringhurst (1951) in
California, also noted that ideally temperatures should be above 6.5/19°C and 10/20°C
(min./max.) for cultivars with Group A and B floral behaviour, respectively . Rainfall in the areas

17
of origin was mostly moderate to high, from about 650 mm (at high altitudes) to over 1500 mm
(Praloran, 1970). Samson (1986) suggested that 1800 mm is a likely optimum for tropical
(lowland) conditions, but noted the absence of data. Discussing the responses of avocado to
rainfall and humidity, Gaillard and Godefroy (1995) state that the Mexican race has a relatively
low water requirement, while Guatemalan and West Indian races have an average water
requirement.

2.1.3. Global Avocado Production

2.1.3.1. World avocado production


World production of avocados has increased more than fourfold over the past four decades,
according to FAO. World production in 2010 production was about 3,581,711 metric tons (mt) it
increased to 4,188,912mt in 2012. Although the origin of this crop is Central America, avocado
is nowadays widely cultivated throughout the world (Alcaraz and Hormaza, 2007; Alcaraz et al.,
2011). The world’s leading producers are Mexico (31.1%), Dominican Republic (8.2%),
Colombia (6.4%), Peru (6.1%), and Indonesia (5.9%). Considering these numbers, it is evident
that the American continent conquers the avocado production (70.3%), being also the continent
with the highest amount of harvested hectares (ha), followed by Africa (15.2%), Asia (10.9%),
Europe (1.9%), and Oceania (1.6%) (Statistics Division of the Food and Agriculture
Organization for the United Nations, 2013). The main countries producing avocado fruit within
each continent, where it has been illustrated the fact that Kenya, Mexico, Indonesia, Spain and
Australia are those with remarkably higher production. As this fruit has distinctive and pleasant
sensory attributes, and it is perceived by buyers as beneficial to health, its demand has underwent
an increase over the past few years, a fact that has consequently caused an increase of the area
harvested (Rodrı´guez-Fragoso et al., 2011). In 2013, the world production of avocado was of
about 4.7 million tonnes, increasing 5% over the previous year (Statistics Division of the Food
and Agriculture Organization for the United Nations, 2013). An important agricultural parameter
is the crop yield, which refers to the measure of the production of a specific crop per unit area of
land cultivation. The higher the crop yields, the better the land quality or the higher the
cultivation efficiency. The crop yield for avocado fruit is, globally, of about 9.5 t/ha. The
principal avocado producers present higher values; around 9.5_13.8 t/ha for Mexico, Colombia,
Peru and Indonesia, and 30.0 t/ha for Dominican Republic.

18
A slight increase of the crop yields would generate a considerable impact on avocado production.

Table 2.1. Avocado production of the top 12 producing countries in 2000 and 2008
(FAOSTATS, 2010a)

2.1.3.2. Avocado production in Africa


Over a period of 2005 – 2012 avocado production in Africa has grown but unevenly from
497,339mt in 2005 to 751,881mt in 2012. The leading producers with their volumes in mt in
2012 were Kenya (186,292mt), Rwanda (145,000mt), South Africa (91,603mt), Cameroon
(72,000mt) and DRC (70,000mt). With an exception of South Africa all the other four top
ranking producers lie closer to the equator with tropical conditions. Africa’s annual production
growth has been closer to the global rate that between 2005 and 2012 averaged at 6.4%. Year-on-
year growths have been the highest in Morocco (23%), Tanzania (20%), Rwanda (18%) and
Kenya (10%). In absolute terms Kenya and Rwanda carry more weight due to the larger base
while a country like Tanzania had a smaller base of 22,000mt. It is promising to know that the
crop has market foothold in both export and domestic markets, data from the South African
Subtropical Growers Association indicate that 45 % of total production is exported, 15 % is
processed, 25 % is consumed in local markets and 15 % is sold on the informal markets. While
South Africa ranks low in terms of volume of fruit produced, it has the most advanced subsector

19
and is among the top five of exporters. Other Africa exporters are Ethiopia, Cameroon, Rwanda
and Kenya. It also has a well advanced R&D base.
Table 2.1 Africa top 10 avocado producing countries (2008-2012, in mt) Faostat 2014

2.1.3.3. Avocado production in Ethiopia


Fruits have significant importance with a potential for domestic and export markets and
industrial processing in Ethiopia. The main fruits produced and exported are banana, citrus fruits,
mango, avocado, papaya and grape fruits (Zeberga, 2010). Owing to its shortest introduction to
Ethiopia, these days the crop is produced in several countries where Ethiopia stands the 10th
leading producer and 6th most important consumer in the world (FAOSTAT, 2010).
Avocado was first introduced to Ethiopia in 1938 by private orchardists in Hirna and Wondo-
genet and production gradually spread into the countryside where the crop was adapted to
different agro-ecologies (Edossa, 1997). Annual avocado production in Ethiopia is 25633.16
tons. The crop is now produced by 1,149,074.00 farmers countrywide who collectively farm
more than 8938.24 ha of land (CSA, 2012/13). According to Garedew (2010) even though
avocado has economically and socially play a significant role its production is confronted by a
number of constraints;- this are degeneration of fruits, disease problem and absence of
agronomic practices.
According to FAO, (2010) description of some varieties introduced in Ethiopia and presently
available includes: Hass: high yielding, resistant to main pests and diseases. not presenting a

20
marked biennial fruiting behavior. Fruit size variable; oil % in the fruit: medium, month to ripen:
9, seed size: small; cold tolerance: medium. Pinkerton: high yielding; fruit size; medium; oil %
in the fruit: high; month to ripen: 6-8; seed size: big; cold tolerance: medium. Fuerte: a Mexican
Guatemalan cross; medium yielding, fruit size , small to medium; oil % is high; month to ripen
5-6; seed size : tolerant to frost. Bacon: high yielding; medium size fruit; oil % high; tolerant to
cold -5oC. Ettinger: a Mexican Guatemalan cross, resistant to cold Nabal: Guatemalan type, big
size fruit; suitable for warm climate. In general, fruit production is still backward, the business is
underdeveloped and the private sector is not much attracted. In connection with this lack of
access to improved varieties, production is exclusively based on distribution of mixed materials;
consequently the local seed system has come out as best-bet arena and is now a common route
for seedling dissemination (Ayelech, 2011 ). CSA (2013) indicated Avocado as one of the
second potential fruit crop produced in Ethiopia.
Ethiopian Farmers produce avocado without knowing its variety. Knowing their variety is useful
in selecting variety which give high yield, high oil content and also in determining time of
harvesting. In Ethiopia avocado is consumed mostly as a juice either alone or mixed with other
fruit juice and also as a salad. Its juice is consumed round a year especially during fasting days
and mostly utilized by urban dwellers rather than rural dwellers due unavailability of it
everywhere since avocado is not produced all over Ethiopia. Eating small amount of avocado
feels stomach full which incase useful in reducing weight due to its high fat content. Its fat
content among different varieties in Ethiopia is not yet investigated which was determined in this
study.

2.1.4. Avocado fruit Uses and Composition


Avocados have significantly higher lipid content (8–25%) as compared to other fruits. The
avocados have diverse fats yet they are one of the best foods one can eat. They are full of
nutrients and heart-healthy compounds. According to USDA report in 2004, each 100g (3.5oz) of
avocado pulp gives 670KJ (160Kcal) of energy; 75% of which is from its fat. It contains 2.13g
saturated fatty acid, 9.80g monounsaturated and 1.82g polyunsaturated fatty acids. 2g of that
amount was protein while water was 73.23g. The avocado also contains many vitamins and
minerals; especially it contains 35% more potassium than banana which has 358mg per 100g.
75% of the high fibre content is insoluble while 25% is soluble (Naveh et al., 2002). The health
benefits of avocados are quite obvious, from their nutritional components and hence cannot be

21
overemphasized. High intake of avocados lowers blood cholesterol levels (low density
lipoprotein, harmful cholesterol) due to its high content of High density lipoproteins (HDL),
helpful cholesterol (Naveh et al., 2002).
The avocados are a great source of liteine, a carotenoid that works as an antioxidant and helps
protect against eye disease. Avocado helps in weight loss because the monounsaturated fats
make one feel full and resist temptation to eat. It contains good amount of fiber both soluble and
insoluble. Fiber is needed by the digestive system to run smoothly. Oleic acid is a fat that 20

22
activates the part of the brain that makes one feel full, and it is also present in avocado. Oleic
acid in avocado has been shown to produce greater level of satiety than less healthy saturated fats
and Trans fats contained in processed food. The processed products of avocado pulp include the
paste, puree, and guacamole. Guacamole is a fruit pulp seasoned with salt, onion, lemon, pepper
and tomato, being produced not only in an artisanal way but also marketed by some US
companies (Daiuto et al., 2011).
The sensory quality of guacamole of Hass variety made without chemical additives and stored
under refrigeration was evaluated according to the type of packaging used. A greater consumers’
acceptance was observed for the product stored in container with gas barrier when compared to
that stored in polyethylene package (Daiuto et al., 2011). Although these authors have also
considered that the heat treatment may have been effective on the polyphenol oxidase
inactivation, it can result in the development of bitterness and off-flavors in avocado, which
changes the guacamole texture, negatively contributing to a mashed appearance.
Chaves et al. (2013) studied avocado pulp Margarida variety dehydrated and defatted by cold
pressing and avocado oil to partially replace wheat flour and butter, respectively, in whole grain
crackers. The authors reported that the flour from avocado pulp, in general, showed
characteristics similar to those of conventional flour and whole wheat flour. The biscuits had
higher minerals and fiber levels, with good sensory acceptance.
Meat derivatives can also be supplemented with avocado pulp, since most of these processed
foods contain relatively high levels of saturated fats in the formulation whose consumption is
restricted by health issues. Thus, an alternative to reduce and enhance fatty acids balance is the
incorporation of fats or vegetable oils in emulsified meat products. The replacement of animal
fats by vegetable oils in meat products has been studied with positive effects on the chemical,
physical and sensory characteristics of the products, but with negative effects on water activity
and texture (Lugo, 2006).

2.2. Avocado fruit tree and canopy management


Canopy management is the manipulation of tree canopies to optimize the production of quality
fruits. The canopy management, particularly its components like tree training and pruning,
affects the quantity of sunlight intercepted by trees, as tree shape determines the presentation of
leaf area to incoming radiation. An ideal training strategy centers around the arrangement of
plant parts, especially, to develop a better plant architecture that optimizes the utilization of
23
sunlight and promotes productivity. The basic problem with overcrowded orchards is insufficient
light (Stadler and Stassen, 1985). There are several systems to manage tree size and improve
light interception and penetration, including selective limb removal (individual limbs are
removed to maintain tree size and inter-row access); mechanical pruning (trees are pruned to
form a hedgerow); stag-horning (trees are pruned back to a stump and allowed to re-grow); tree
thinning (alternate rows or trees within a row are removed as orchards begin to crowd) and tree
removal (whole blocks removed after 10-15 years and replaced with new trees). With avocado
there remains opportunity and challenges to reduce the time between planting and full canopy
development as well as maintain a productive orchard once an effective full canopy has been
attained. Orchard planting densities varying from about 90 to 1600 trees ha−1, with trial
plantings in Chile of up to 2500 trees ha−1, emphasize uncertainty in orchard design, aggravated
by big differences in climate and soil. Recent studies of the light regime in an avocado canopy
(Hadari, 2004; Heath et al., 2005) emphasize the importance of canopy light exposure
optimization over the life of the orchard.

2.3 Avocado fruit tree light interception and light use efficiency

2.3.1 Light interception and photosynthesis of avocado fruit tree canopy


Avocado canopies consist of relatively long lived leaves that are exposed to varying light
conditions throughout their lifespan. Because of the flushing nature of avocado shoots, most
leaves will develop in full sun then exist in deep shade after re-growth has occurred, while other
leaves develop in and remain in shaded portions of the canopy. Leaves that develop in shade
have lower chlorophyll a/b ratios and lower carotenoid concentrations (Förster et al., 2009).
Shade leaves of avocado have lower photosynthetic rates not due to differences in stomatal
density (Mickelbart et al., 2000), but due to reduced stomatal conductance at lower light levels
(Sterne et al., 1977).The connection between increased light interception and yield has been
studied intensively, especially in apple orchards. Dry matter productions as well as various
parameters dealing with quality and quantity of fruits were reported to have an empirical
relationship with the amount of light falling on the orchard (Barritt, 1989; Barritt et al., 1991;
Wunsche et al., 1996). However, the agreement was better for the primary biomass production
than for the fruit yield, due to the additional parameters affecting the yield (Palmer, 1999). Full
canopies of orchard crops intercept only 65% to 70% of the available radiant energy, thereby

24
creating an upper limit to the production potential (Jackson, 1980). It appears that maximum
photosynthetic rate occurs when leaves are exposed to at least 0.3 of the full sunlight intensity
(Heinecke, 1966). The fraction of the total sunlight intensity mentioned above is evidently not
always adequate for the normal development of vegetative and reproductive buds. Palmer (1977)
found that flower-bud differentiation in apple trees is more sensitive to shading than vegetative
growth. Heinecke (1966) reported insufficient coloring of apples when the light intensity was
below 0.4 and when it was lower than 0.5 fruit size was adversely affected. Jackson (1978)
confirms that high light interception is a prerequisite for maximal yield, whereas shading causes
a reduction in flower-bud formation, fruit size, and fruit color. In Florida, rejuvenation of
crowded Avocado orchards as a result of topping and tree removal, proved to be successful.
Although the orchard was less dense (130 trees ha-1 instead of 276 trees ha-1) and the tree height
was reduced to 4.8m, the yield increased from 6.9 t ha-1 to 19.8 t ha-1 (Crane et al., 1992). The
result from this experiment emphasized the great importance of light interception, since it is the
main reason for the increase in yield. Reports from South African Avocado pruning trials
(Stassen et al., 1999) showed improvement in the yield from pruned hedgerows. The yield
response to pyramid shaping and lowering of the tree height was considerable. Both practices
aimed at increasing the light intercepted by the hedgerow. The pyramidal shape improved
canopy-sun grazing angle and the height lowering prevented inter-row shading. In a pruning trial
aimed at testing the effect of lowering the height of Avocado trees (Thorp and Stowell, 2001), 6
trees were pruned down to 4 and 6 meters and fruit yield was collected from various heights.
Reducing tree height resulted in a shift in the location of most of the Avocado fruits. The same
amount of fruit previously found in the 4-6 m height layer in the 6m high trees was found in the
2-4 m height layer in trees pruned to 4 m height. In a research done in New Zealand the harvest
quality of fruit exposed to sun was compared with that of completely shaded fruit. Significant
differences were found between the two fruit types and between the different sides of fruits
exposed to sun. Namely, fruits exposed to sun had higher dry matter and higher levels of
potassium, calcium, magnesium and oil (Woolf et al., 1999).

2.3.2. The Avocado fruit trees response to light


Due to its rain forest origin, characterized by competition for light, the Avocado (Persea
americana Mill.) tree have a natural vegetative bias towards greater allocation of
photoassimilates to shoot growth than to reproductive organs (Schaffer and whiley, 2002). This

25
vegetative bias results in rapid production of short-lived leaves and increased shading of the
canopy, thus reducing the number of well-lit terminal shoots capable of flowering (Schaffer and
Whiley, 2003). The growth of Avocado reproductive organs consumes large amounts of energy,
which is then being stored. Indeed, CV. Fuerte fruits with 17% oil content and a flesh-to-seed
ratio of 4:1 (fresh mass) store Energy equivalent value of 8 GJ ton-1 compared to 2.9 GJ ton-1
for Valencia oranges and 2.6 GJ ton-1 for apples. The leaves can also store large amounts of
carbohydrates and minerals, with episodic growth flushes resulting in leaves of varying age and
photosynthetic efficiency (Wolstenholme, 1987). It is generally agreed that the low yielding
potential of Avocado orchards is due to two main factors: the high oil content of the fruit (oil is
2-3 times more energy-expensive than carbohydrate) and the large seed with its concentrated
food reserves (Wolstenholme, 1987).

2.4 Avocado tree flower, pollination mode and pollinizers

2.4.1. Avocado tree flower and Flowering

2.4.1.1. Avocado tree flower


The avocado flower is circular, about 1 cm in diameter. It is bisexual, having both female and
male reproductive organs. The flower consists of one pistil and six trimerous alternate whorls:
two whorls of greenish-yellow tepals, three of stamens and one of short arrow-shaped
staminodes . Each of the nine stamens bears an anther with four pollen sacs. The valves of the
pollen sacs turn up while opening, and draw out a pack of pollen grains attached to them. The
avocado pollen grain is spherical and is covered with numerous conical spinules. A pair of oval,
yellow- orange nectaries is located at the base of each of the three inner-stamen filaments. The
three yellow-orange staminodes also function as nectaries. The pistil is located centrally, and
consists of a greenish, ball-shaped, superior ovary, a hairy, slender, cylindrical style and a cone-
shaped stigma, 0.3 to 0.6 mm in diameter. The stigmatic surface is composed of elongated
papilla cells. Usually, it is somewhat depressed in its center. All cultivars have a similar flower
structure, though they may differ slightly with respect to flower size and some other features
(Nirody, 1922; Stout, 1933; Bergh, 1969; McGregor, 1976; Davenport, 1986; Ish-Am and
Eisikowitch, 1991b). The flowers are carried on terminal panicles. A typical panicle carries a few
hundred to more than a thousand flowers. New flowers open daily during the long flowering
period. Each flower opens twice (dianthesis), usually on two successive days, in a dichogamous-

26
protogynous rhythm . During the first female (pistillate) opening (Photo 1), the stigma is exposed
and white. The stamens with the closed valves are parallel and close to the tepals and nectar is
secreted by the three staminodes. At the beginning of the second male (staminate) opening the
tepals are 10% to 20% longer. The three inner stamens move towards the style, and their
filaments lengthen until their anthers more or less cover the stigma. The six outer stamens also
lengthen, move up and hold a position of about 45° to the pistil. Anther dehiscence takes place
about 1 to 2 h later, first by the two lower valves of each anther, and about 1 h later by the two
upper valves. Nectar is secreted by the six nectaries, while the three staminodes move closer to
the ovary and turn brown. The stigma gradually shrinks, and also turning brown (Nirody, 1922;
Stout, 1933; Bergh, 1969; McGregor, 1976; Davenport, 1986; Ish-Am and Eisikowitch, 1991b).
During anthesis, flower shape changes gradually in a regular sequence, which was divided by
Ish-Am and Eisikowitch (1991b) into ten distinct morphological stages. These flower stages are
similar in all cultivars, and are somewhat influenced by the weather. On hot and dry days, the
male flower stigma and staminodes dry upon anthesis, and the tepals bend towards the pedicel.
In cool weather, however, both stigma and staminodes of the male-stage flower stay fresh during
the first dehiscence stages and the tepals spread upright to the style, or even remain only partially
open. Under cool conditions, some cultivars open their female flower only partially or not at all
(Nirody, 1922; Stout, 1933; Bergh, 1969; Ish-Am and Eisikowitch, 1991b).

2.4.1.2 Avocado tree flowering


All avocados display unique flowering behaviour, which may be termed ‘diurnally synchronous
dichogamous protogyny, with intermediate closing’. The flower opens twice, first as a female
and then as a male. Each flower-stage opening and closing occurs nearly synchronously within
the tree (and the cultivar). Based on flowering rhythm, all avocado cultivars are divided into two
complementary flowering groups: In warm weather, ‘Group A’ cultivars open female-stage
flowers from the morning till noon, and male-stage flowers during the afternoon. ‘Group B’
cultivars, on the other hand, open female-stage flowers in the afternoon and male-stage flowers
during the morning hours. This flowering rhythm may be termed ‘temporal dioecy’ (Clark, 1923;
Stout, 1923, 1933; Robinson and Savage, 1926; Bergh, 1969; McGregor, 1976; Papademetriou,
1976b; Davenport, 1986; Ish-Am and Eisikowitch, 1991b).
Many avocado cultivars also present a daily phase of overlap within the tree, of coinciding
dehisced male-stage and open female-stage flowers, which usually takes place for a period of 1

27
to 3 h. In some cultivars this overlap period is almost constant, regardless of temperature, but in
others it gets shorter during warmer weather and may disappear on hot days. In cool weather,
there is a significant delay in the whole daily flowering sequence, which may result in complete
reversal of the times of day when female-and male-stage flowers are open (Clark, 1923; Stout,
1923, 1933; Robinson and Savage, 1926; Lesley and Bringhurst, 1951; Gustafson and Bergh,
1966; Bergh, 1969; McGregor, 1976; Papademetriou, 1976b; Sedgley, 1977; Sedgley and
Annells, 1981; Sedgley and Grant, 1983; Davenport, 1986; Ish-Am and Eisikowitch, 1991b).
Although the beginning (and the end) of each flower-stage manifestation is synchronized within
the tree and among trees of a cultivar, the individual flowers do not open and close
simultaneously. Rather, they proceed through each stage individually, over a period of 2 to 3 h.
Flowers that open earlier also proceed earlier to the following stages. Therefore, during most of
the flowering day, several consecutive flower stages occur concurrently within the panicle (Ish-
Am and Eisikowitch, 1991b).

2.4.2. Avocado tree pollination Modes


2.4.2.1. Cross-pollination
Cross-pollination occurs between group B male-stage and group A female-stage flowers in the
morning (in warm weather), and vice versa in the afternoon. It may also occur among different
cultivars of the same flowering group, when there is a period of overlap between female and
dehiscing male openings (Stout, 1923, 1933; Robinson and Savage, 1926; Bergh, 1969;
Davenport, 1986; Ish-Am and Eisikowitch, 1991a,b). Cross-pollination effectiveness depends on
the distance between the ‘pollenizer’ (pollen donor) and the pollinated trees, on the effective
overlap period between female and male openings, and on pollinator mobility, density and
effectiveness. In many cases, cross-pollination of a group A cultivar by a group B one is more
efficient than the other way around, due to the greater overlap period between group A female
and group B male flowering relative to that between the opposite blooms (Stout, 1933; Ish-Am,
1994; Papademetriou, 1976b; Ish-Am and Eisikowitch, 1991b).

2.4.2.2. Close-pollination
Close-pollination occurs during the phase of female- and dehisced male-stage flower overlap
within the tree (Stout, 1923, 1933; Robinson and Savage, 1926; Lesley and Bringhurst, 1951;
Bergh, 1969; Snir, 1971; Papademetriou, 1976b; Davenport, 1986; Ish-Am and Eisikowitch,

28
1991a,b). Since close-pollination takes place when male- and female-stage flowers are in close
proximity, its efficiency may be very high. It depends on both length and effectiveness of the
overlap period within the tree, and on pollinator density and effectiveness. Most Mexican- and
Guatemalan-type cultivars, and their hybrids, present a daily effective period of overlap within
the tree. Generally, close-pollination is more efficient in group A cultivars, since their open
female flowers overlap with young pollen-releasing male flowers, and less so in group B types,
where newly opened female flowers overlap with old male flowers, which have finished pollen
release and are closing (Ish-Am and Eisikowitch, 1991b; Ish-Am, 1994).

2.4.2.3. Self-pollination
Self-pollination occurs when pollen released at the male stage, reaches the stigma within the
same flower. This process does not necessarily demand pollinator involvement, and may be
facilitated by wind or gravity. Self-pollination of the male flower is a common phenomenon, but
in most cases does not lead to fertilization (Stout, 1923, 1933; Robinson and Savage, 1926;
Bergh, 1969; Snir, 1971; Davenport, 1986; Ish-Am and Eisikowitch, 1991a,b). However,
Davenport (Davenport, 1985, 1989; Davenport et al., 1994) concluded that in south Florida,
spontaneous self-pollination is the predominant means for fruit set.

2.4.3. Avocado tree pollinizer


Various pollinating agents visit the avocado flowers for nectar and pollen. These include the
honey bee, various species of wild bees, wasps, flies, and hummingbirds (Chapman 1964). The
consensus of various research workers who have studied the flowering and fruiting of the
avocado is that only honey bees are sufficiently abundant on the blossoms at all times to set
satisfactory crops of fruit (Clark 1923,1924; Clark and Clark 1926; Boyden 1930; Traub et al.
1941; Lemmerts 1942; Lesley and Bringhurst 1951; Winslow and Enderud 1955; Lecomte 1961;
Popenoe 1963). Many observers have noted that a bee tends to visit a single tree and thus fails to
afford the cross-pollination desired. This can occur when the trees are separated by some
distance, for example, when they are small or spaced too far apart (Bergh 1966). It also occurs
when there is an insufficiency of bees in relation to the number of blooms available. When the
flowers per bee ratio is low, the bees are required to visit many flowers to obtain a load of food
and their efficiency as cross-pollinating agents is increased. Ruehle (1958) stated that good crops
are set consistently in groves a considerable distance from any bee hives hut that the presence of

29
trees would increase production. Wolfe et al., (1942, 1946) stated that it is quite possible that a
hive of bees per acre with sets of five in the middle of each 5-acre tract would materially increase
production. Popenoe (1963) stated that honey bees are probably necessary for good pollination
unless there is an abundance of wild bees in the area. In an excellent survey of the reasons for
low yield of avocados in California, Bergh (1967) unequivocally stated: "Practically every
avocado fruit set means that a honey bee transferred pollen to that flower from some other
flower. Gravity or wind may act, but they are so rare they can be ignored by the practical
avocado grower." Further on, he stated, "At the present time the California avocado industry is
dependent upon the honey bee. The greater the bee population, the more likely the bees are to
travel from flower to flower and so make the best of such inter-flower overlap in male and
female stages as may be present. This is probably the chief source of avocado set in California."

2.5. Avocado fruit orchard and Honey bee population


The honeybee (Apis mellifera L.) is apparently the major pollinator of avocado in all countries.
Honeybees visiting avocado flowers usually walk among neighbouring flowers, and fly between
inflorescences. While visiting the relatively small flower the bee often slips, and grasps other
parts of the inflorescence (Davenport, 1986; Vithanage, 1990; Ish-Am and Eisikowitch, 1991a,
1993). Honeybees usually collect avocado nectar, or nectar with pollen, and rarely only pollen.
The nectar-collecting bees, as well as the nectar-with-pollen collectors, visit both female- and
male-stage flowers, and as such may serve as effective pollinators. In contrast, the pollen-only-
collecting bees visit almost exclusively male flowers, and do not contribute to pollination (Stout,
1933; McGregor, 1976; Davenport, 1986; Free, 1993; Ish-Am and Eisikowitch, 1993). While
visiting a dehisced male flower, the bee’s body becomes dusted with pollen, which the bee
cleans off after every two to four visits, whilst hovering or hanging on a leaf. The nectar-and-
pollen collectors pack the pollen into their curbiculae, forming pellets, whereas the nectar-only
collectors ‘deliberately’ unload the pollen and throw it down (Ish-Am and Eisikowitch, 1993).
The pollen-only collectors’ visits are very short: in about 1 s they touch the anthers while
hovering, or while landing for an instant. These bees may also perform one to two refueling
nectar collections per ten pollen collections, during which they may visit female flowers, if
present in the vicinity (Ish-Am and Eisikowitch, 1993). The positions of bees visiting female-
and male-stage flowers of equivalent form are very similar . Only limited and defined zones of
the bee’s body, the ‘pollinating zones’ contact the flower’s reproductive organs. While visiting
30
the male flower, the bee touches the exposed pollen on the open valves with its vertex,
proboscidal fossa, legs and some ventral regions collecting large amounts of pollen there. The
same ‘pollinating zones’ also touch the stigma of the female flower due to the similarity of the
female- and male-stage flowers, and to the similar location of the stigma and inner stamens’
anthers (Ish-Am and Eisikowitch, 1993).

Most honeybees that visit avocado during the period of overlap between female and male
openings within a tree, move between the male and female flowers, collect nectar and pollen, or
nectar only, and efficiently carry out close-pollination. (Clark, 1923; Gustafson and Bergh, 1966;
Gazit, 1976; Davenport, 1986; Ish-Am and Eisikowitch, 1993).

For cross-pollination implementation, honeybees need to move between pollen-releasing male-


flower trees and female-flower trees of a different cultivar. In mature avocado orchards, an
average 40% of the bees were found to move between adjacent trees in 10 min (Ish-Am and
Eisikowitch, 1998b). However, this constitutes short-range movement, since during foraging
most honeybees visit only one to three adjacent trees (Clark, 1923; Stout, 1923, 1933; Free and
Spencer-Booth, 1964; Bergh et al., 1966; Gustafson and Bergh, 1966; Bergh, 1967; McGregor,
1976; Davenport, 1986; Visscher and Sherman, 1998; Vithanage, 1990; Free, 1993; Hofshi,
1995, 2000). Indeed, scout bees move, while foraging, farther throughout the orchard; but, under
avocado orchard conditions they were found to comprise only 2 to 4% of the field bees (Stout,
1933; Ish-Am and Eisikowitch, 1998b). Therefore, honeybees serve as efficient cross-pollinators
only in the case of neighbouring trees with complementary flower types, and their efficiency
decreases sharply with increasing distance between the pollen source and the female bloom (Ish-
Am, 1994; Ish-Am and Eisikowitch, 1998b).

The attractiveness of the avocado bloom to honeybees under Mediterranean climatic conditions
appears to be low compared to numerous species that are in bloom at the same time, such as
citrus, litchi and wildflower species. Therefore, honeybee foragers from hives placed in the
avocado orchard often abandon the avocado and collect pollen and nectar from competing
blooms. In Israel, this phenomenon presents a major yield-limiting factor for the early- and
medium-blooming cultivars, which flower in March-April, during the blooming season of citrus
and many wildflower species (Clark, 1923; Stout, 1923, 1933; Gustafson and Bergh, 1966;
Bergh, 1967; Gazit, 1976; McGregor, 1976; Papademetriou, 1976b; Tzfati, 1981; Eisikowitch

31
and Melamud, 1982; Davenport, 1986; Shoval, 1987; Visscher and Sherman, 1998; Vithanage,
1990; Ish-Am and Eisikowitch, 1998a; Hofshi, 2000).
This low attractiveness to honeybees has been attributed to the avocado flower and flowering
properties, which are not well-suited to this pollinator (Faegri and Pijl, 1979; Kevan and Baker,
1983; Vithanage, 1990; Ish-Am and Eisikowitch, 1993; Visscher and Sherman, 1998; Ish-Am et
al., 1999). The avocado flower is greenish-yellow, has a slightly bitter smell and its nectar is
fully exposed. It has radial symmetry, lacks a landing platform and nectar trails, and is somewhat
small for the honeybee, while the inflorescence is too sparse to be visited as a unit (Davenport,
1986; Vithanage, 1990; Ish-Am and Eisikowitch, 1993). Moreover, neither avocado pollen nor
its nectar suits fully the honeybee’s needs (Ish-Am, 1994

3. MATERIAL AND METHODS

3.1. Description of the Study Area


The study will be conducted at Sidama Regional State Teferi kela District which is located in
the southern Ethiopia about 63 km (39 miles) from Hawassa, the capital city of Sidama Regional
State and 284 km (176 miles) from the capital Addis Ababa. According to the classification used
in Ethiopia, the climatic condition of the district is characterized as Woina Dega (Mid-altitude)
Zone. The area receives a bimodal rainfall where the high rains are between March to April and
the small rains are from September to October. The major cultivated flowering fruit crop in the
district includes perennial crops including Coffee, Avocado and Mango (Agricultural and
Natural Resource Management Office, 2020). It is found at latitude of 6.50029820 and longitude
38.4000. It has an Altitude (meters), Lat (DMS), 6° 30' 0N, and Long (DMS), 38° 23' 60E. Its
geographical coordinates are 6° 30' 0" North, 38° 24' 0" East and its original name (with
diacritics) is Teferī Kēla. The soil of the experimental site is mainly clay loam. Mean annual
rainfall of the area 1600- 1800 mm, with minimum and maximum temperature of 18.5℃- 32℃ .

3.2. Experimental Design and Treatments, procedure


The experiment will be carried out in factorial randomize complete block design (RCBD) with
three replication. The avocado fruit trees will select depend on their size and with different

32
heights at study area. The factor A treatments consist of 1) control 2) prune unproductive
branches. Measurements of light reaching the inside of the avocado canopy will states as
percentages relative to the amount of light above the canopy and photosynthetically active
radiation (PAR) data will be collected. The factor B treatments consist of 1) control without hives
2) four hives/ha, 3) six hives/ha and the honey bee hives (A.m. scutellata) top-bar type will be
used and located in the center of orchard and within this orchard three sites will be selected,
covering a range of distance to the hives: 30, 60 and 90 m . At each site four trees will samples, with
a total of 12 trees in orchard.

3.3. Data collection


Data will be collected using the standard procedures and following physiological, light quality,
fruit quality and yield parameters

 Canopy height (m)


The canopy height will measure with a good accuracy using small footprint lidars. This is
essentially accomplished by subtracting the last return altitude (ground) from the
corresponding first return altitude (canopy surface).
 Canopy volume (m3/ha)
Canopy volume is generally calculated using the following predefined volume formula .
Canopy Volume = Canopy Height *(Crown Diameter) 2 * Multiplier
 Leaf Area Index (LAI)

The Leaf area index is a dimensionless quantity that characterizes plant canopies. It is defined as


the one-sided green leaf area per unit ground surface area.

LAI = leaf area / ground area, m2 / m2

 Photosynthetically active radiation (PAR)


The PAR will measure once during the day under diffuse light conditions using an AccuPAR ceptometer
(Decagon Devices Inc.).
 Red light,
 Far- red
 Red to fared ratio.
 Density of bees per tree (BPT)

33
The BPT will record during the flowering period and quantify by a person walking around a tree and
counting with a manual counter the presence of the bees per minute.
 Pollination rate and efficiency (PR and PE)
A sample of stigmas per tree will be collected in the female flowering stage. The pollination rate
(percentage of stigmas pollinated \ total stigmas collected) and pollination efficiency (average
number of pollen grains in the stigma, only for pollinated stigmas) will quantify for each treatment.
 Percentage of fruit set initial (PFSi)
In order to determine the PFSi, the number of flowers (flowering) and the number of fruits (four
weeks after the end of flowering) will record on five inflorescences per randomly selected
tree.                        PFSi= Number of fruits set / Number of flowers * 100.
 Percentage of fruit set final (PFSf)
In order to determine the PFSf, the number of flowers (flowering) and the number of final formed
fruits will be record on five inflorescences per randomly selected tree.
PSFf = Number of fruits formed / Number of flowers * 100.
 Total number of fruits per tree (NFTT)
The NFTT will record after the end of flowering and all the fruits of each tree will be counted.
The physical quality parameters of avocados include skin color, firmness, texture, and physiological
disorders, Oil content, Dry matter

3.4 Statistical Analysis


The Duncan test will be used for the comparison of means for each factor. Subsequently, a
Pearson correlation analysis will perform on the evaluate variables. Data will analyzes by
ANOVA using (SAS) statistical analysis software, with significant differences among means
determined by Least Significant Difference (LSD) at p≤0.05.

The collected data will be subjected to Analysis of Variance (ANOVA) using the Statistical
Analysis System (SAS) Software version (9.0). Mean separation between treatments will done
using Least Significance Difference (LSD) at 5% and 1% probability base on the ANOVA
results a indicated by Gomez and Gomez (1984).

34
4. WORK PLAN
All the field activities and data collection will be accomplished starting from October 2021 G.C 1st
week up to end of January 2022 G.C. Data entry and write will be completed on February 2022
G.C. The detailed activity schedule is listed below (Table1).

Table 4.1: Work plan schedule


s/no Activities to be done 2021 G.C 2022 G.C
Aug Sept Oct Dec Nov Jan Feb March April May June July
1 Problem
identification

2 Proposal 

preparation

3 Proposal defence 

4 Site selection 

5 Lay -out 

6 Data collection  

7 Data analysis 

35
8 Thesis write up and 

edition

9 Thesis first draft 

submission

10 Final thesis 

defence

5. BUDGET BREAKDOWN
Table 5.1: Stationery

S No Item Unit Amount Cost


Unit price Total price birr
birr
1 Paper A4 size Rim 4 200 8,00
2 Pen No 20 10 2,00
3 Pencil No 20 1 20
4 Eraser No 4 8 32
5 Computer writing No 200 5 1,000
6 Photocopy No 200 1 200
Sub Total 2,252.00

Table 5.2: Transport Cost

S No Item Unit Amount Cost


Unit price Total price birr
birr
1 Fuel Lit 1000 25 25000
2 Motor oil and lubricants Lit 10 120 1200
Sub total No 26,200

36
Table 5.3: Farm and laboratory inputs

S No Item Unit Amount Cost


Unit price Total price

1 Kenya Top Bar Bee No 20 2000 40,000


Hive
2 Shade net No 70 1000 70000
3 Land Compensation Hec 3 3500 10,500
4 Light measurement Various
Sub total

Table 5.4: Labor cost

S No Items Units Cost


Days laborer PPD birr Total
1 Site preparation MPD 5 60 100 30,000
2 Bee transferring MPD 5 20 100 10,000

3 Shed construction Various 10,000


Beekeepers protective 2000
clothes
Sub total

Table 5.5: Personnel cost

S No Items No of days Rate Total

1 Researcher
2 Advisors
3 Driver
Sub Total

37
6. REFERENCE
Alcaraz, M.L., Hormaza, J.I., 2007. Molecular characterization and genetic diversity in an
avocado collection of cultivars and local Spanish genotypes using SSRs. Hereditas. 144,
244_253.
Alcaraz, M.L., Montserrat, M., Hormaza, J.I., 2011. In vitro pollen germination in avocado
(Persea americana Mill.): optimization of the method and effect of temperature. Sci.
Hortic. 130, 152_156.
Avilán, L., & Rodríguez, M. (1995). Época de floración y cosecha del aguacate en la región
norte de Venezuela. Agronomía Tropical, 45, 35-50.

Avilán, L., & Rodríguez, M. (1995). Época de floración y cosecha del aguacate en la región
norte de Venezuela. Agronomía Tropical, 45, 35-50.
Ayelech, T. (2011). Market chain analysis of fruits for Goma woreda Jimma zone, Oromia
regional state’ A Thesis Submitted to School of Graduate Studies of Haramaya
University.
Barritt, B. 1989. Influence of orchard system on canopy development, light interception and
production of third year granny apple trees. Acta Horticulturae 243:121-130.
Barritt, B., C. Rom, J. Konishi, and M. Dilley. 1991. Light level influences spur quality and
canopy development and light interception influences fruit prod
Bastias, R. & Corelli-Grappadelli, L. (2012). Light quality management in fruit orchards:
Physiological and technological aspects, Chilean Journal of Agricultural Research, vol.
72, no. 4, pp. 574-581.
BERGH, B. O. 1992. The origin, nature and genetic improvement of avocado. Cal. Avoc. Soc.
Yrbk. 76: 61–75.
BERGH, B. O. 1995. Avocado: Persea americana (Lauraceae). In J. SMARTT AND N. W.
SIMMONDS, EDS., Evolution of Crop Plants. Longman Scientific & Technical, Essex,
UK

38
Bergh, B.O. (1977). Factors affecting avocado fruitfulness. In Saults, J. Philips, R y Jackson, L.
(eds.) Proceedings of the First International Tropical Fruit Short Course: The Avocado.
University of Florida; Gainesville, FL, USA; 83-87.
Bergh, B.O. 1968. Cross-pollination increases avocados set. The California Citrograph 53:97-
100.

Bergh, B.O. 1969. Avocado, Persea americana Miller. In: Ferwerda F.P., Wit F. (eds): Outlines
of Perennial Crop Breeding in the Tropics. Veenman & Zonen N.V., Wageningen,
Netherlands pp: 23-51.

Bergh, B.O. 1977. Factors affecting avocado fruitfulness. In: Saults, J.W. Philips, R.L. Jackson,
L.K. (eds.). Proc. First Intl. Trop. Fruit Short Course: the Avocado. Univ. of Florida,
Gainesville. 83-88.

Bergh, B.O. Garber, M.J. 1964. Avocado yields increased by interplanting different varieties.
California Avocado Society Yearbook 48:78-85.

Bost, J.B., Smith, N.J.H., Crane, J.H., 2013. History, distribution and uses. In: Schaffer,B.,
Wolstenholme, B.N., Whiley, A.W. (Eds.), The Avocado. Botany, Productionand Uses.
CABI, Wallingford, UK, pp. 10–30.

BUCKLER IV, E. S., D. M. PEARSALL, AND T. P. HOLTSFORD. 1998. Climate, plant


ecology, and Central Mexican archaic subsistence. Curr. Anthropol. 39:152–164
Can-Alonzo, C., Quezada-Euán, J., Xiu-Ancona, H., Moo-Valle, H., Valdovinos-Nunez, H.,
Medina-Peralta, S. (2005). Pollination of “criollo” avocados (Persea americana) and the
behavior of associated bees in subtropical Mexico. Journal of Apicultural Research,
44(1), 3-8.
Castañeda, A., Equihua, A., Valdés, J., Barrientos, A., Ish-Am, G., Gazit, S. (1999). Insectos
polinizadores del Aguacate en los estados de Mexico y Michoacan. Revista Chapingo
Serie Horticultura, 5, 129-136. Retrieved from http://bio-nica.info/Biblioteca/Ish-Am
polinizadores Aguacate.pdf
Chanderbali, A.S., Soltis, D.E., Soltis, P.S., Wolstenholme, B.N., 2013. Taxonomy andbotany.
In: Schaffer, B., Wolstenholme, B.N., Whiley, A.W. (Eds.), The Avocado.Botany,
Production and Uses. CABI, Wallingford, UK, pp. 31–50.

39
Chaves, M.A. et al (2013). Preparation of whole cookie using avocado pulp flour and oil.
Boletim do Centro de Pesquisa e Processamento de Alimentos, v.31, p.215-226.
Chen, H., P.L. Morrell, V.E.T.M. Ashworth, M. de la Cruz and M.T. Clegg. 2009. Tracing the
geographic origins of major avocado cultivars. J. Hered. 100:56-65.

Clark, O.I. 1923. Avocado pollination and bees. California Avocado Association Annual Report
8:57-62.

Crane , J. H., B. Schaffer , and T. L. Davenport 1992. Rejuvenation of a mature, non-productive


'lula' and 'booth8' Avocado grove by topping and tree removal. Proc. Fla. State Hort.
Soc. 105:282-285.
CSA, (2013). Agricultural Sample Survey 2012 / 2013; Volume I, Report on Area and
Production of Major Crops, Statistical Bulletin 532; Addis Ababa, Ethiopia.
Daiuto, E.R et al. (2011). Sensory analysis of cold-stored guacamole added with α-tocopherol
and ascorbic acid. Revista Ceres, v.58, n.2, p.140-148.
Davenport, T.L. (1986). Avocado flowering. Horticultural Reviews, 8, 257-289.
Davenport, T.L. Parnitzki, P. Fricke, S. Hughes, M.S. 1994. Evidence and significance of self-
pollination of avocados in Florida. Journal of the American Society of Horticultural
Science 119:1200-1207.

Degani, C. Gazit, S. 1984. Selfed and crossed proportions of avocado progenies produced by
caged pairs of complementary cultivars. HortScience 19:258-260.

Degani, C. Goldring, A. Gazit, S. 1989. Pollen parent effect on outcrossing rate in ‘Hass' and
'Fuerte' avocado plots during fruit development. Journal of the American Society of
Horticultural Science 114:106-111.

Duarte, P.F.,Chaves, M.A.,Borges, C.D., & Mendonca, C.R.B.(2017). Avocado; Characteristics,


health benefits and uses. International News on Fats, Oils and Related materials,
28(3),28-32

Edossa E (1997). Selection of Avocado (Persea americana M.). Collection of Desirable Fruit
Characteristics and Yield at Jimma, Proceedings of the 8th Annual Conference of the
Crop Science Society of Ethiopia, Feb. 26-27, Addis Ababa. Ethiopia. pp. 26-35.

40
Edossa Etissa (1997). Selection of Avocado (Persea Americana) Collection of Desirable Fruit
Characteristics and Yield at Jimma, Proceedings of the 8th Annual Conference of the
Crop Science Society of Ethiopia, Feb. 26-27, Addis Ababa, Ethiopia, pp: 26-35.
Eisenstein, D. Gazit, S. 1989. Effect of self and cross hand pollination on initial fruit set and
fruitlet development in avocado. Alon Hanotea 43:339-348. (In Hebrew).

Evans, L.J., Goodwin, R.M., & Mcbrydie, H.M. (2010). Factors affecting “Hass” avocado
(Persea americana) fruit set in New Zealand. Insect Biology, 63, 214-218.
Faostat. (2014). Production-avocado Ethiopia
http://faostat.fao.org/site/567/DesktopDefault.aspx?PageID=567#ancor
FOREIGN AGRICULTURAL SERVICE, OFFICIAL USDA ESTIMATES. 2006. Fresh
avocados: Production, supply and distribution in selected countries.
http://www.fas.usda.gov/psd/complete_tables/HT P-table6-101.htm (accessed january 11
2007).
Förster, B., Osmond, C.B. and Pogson, B.J. (2009) De novo synthesis and degradation of Lx and
V cycle Pigments during shade and sun acclimation in avocado leaves. Plant Physiology
149, 1179–1195.
Free, J.B. 1993. Insect Pollination of Crops. 2nd ed. Academic Press, 684 p. Gazit, S. 1977.
Pollination and fruit set of Avocado. In: Saults, J.W. Philips, R.L. Jackson, L.K. (eds.).
Proc. First Intl. Trop. Fruit Short Course: the Avocado. Univ. of Florida, Gainesville. 88-
92.

Gaillard, J.P. (1987) L’avocatier: Sa Culture, Ses Produits. Editions Maisonneuve et Larose and
ACCT, 419 pp. (in French.)
Gaillard, J.P. and Godefroy, J. (1995) Avocado. The Tropical Agriculturist Series,
CTA/Macmillan Education, London, 120 pp.
Gazit, S. Degani, C. (2002): Reproductive Biology of the Avocado. In: Whiley, A.W. Schaffer,
B. Wolstenholme, B.N. (eds.): The Avocado: Botany, Production and Uses. CABI
Publishing, Wallingford UK. 101-133. Goldring, A.

Gazit, S. Degani, C. 1987. Isozyme analysis of mature Avocado embryos to determine


outcrossing rate in a 'Hass'plot. Journal of the American Society of Horticultural Science
112:389-392.

41
Gazit, S. Gafni, E. 1986. Effect of hand pollination with different pollen donors on initial fruit
set in avocado. Israel Agrisearch 1:3-17 (In Hebrew, English Abstract).

Gazit, S., & Degani, C. (2002). Reproductive Biology, In The avocado: botany, production and
uses (pp. 101-127). Oxon: CABI Publishing.
Goodwin, M. (2012). Pollination of Crops in Australia and New Zealand. Australia: Rural
Industries Research and Development Corporation.
Hadari, M. (2004) A three-dimensional model of the light regime in an avocado orchard. MSc
thesis, Israel Institute of Technology, 98 pp.
Hadari, M. (2004). A three-dimensional model of the light regime in an avocado orchard. MSc
Thesis, Israel Institute of Technology, Haifa, Israel.
Heath, R., Arpaia, M.L. and Mickelbart, M. (2005) Avocado tree physiology: understanding the
basis of productivity. In: Lovatt, C.J., Holthe, P.A. and Arpaia, M.L. (eds) Proceedings
of the California Avocado Research Symposium. Riverside, California, pp. 87–119.
Heinicke, D. R. 1966. The micro climate of fruit trees. Proc. Amer. Soc. Hort. Sci. 83:1-11.
https:// doi.org/10.26786/ 1920-7603% 28201 2%296
ILRI (2011). Fruits - A synthesis of IPMS value-chain development experiences. ILRI, Nairobi.
Ish-Am, G. (2004). Principios de la polinización del palto – Una breve revisión. In Proceedings
segundo seminario internacional de paltos. (pag. 11) Quillota- Chile.
Ish-Am, G. Barrientos-Priego, A.F. Castañeda-Vildózola, A. Gazit, S. 1999a. Avocado (Persea
americana Mill.) pollinators in its region of origin. Revista Chapingo Serie Horticultura 5
Num. Especial. Proc. of Fourth World Avocado Congress (1999) 137-143.

Ish-Am, G. Eisikowitch, D. 1991a. Possible routes of avocado tree pollination by honeybees.


Acta Horticulturae 288:225-233. Ish-Am, G. Eisikowitch, D. 1991b. New insight into
avocado flowering in relation to its pollination. California Avocado Society Yearbook
75:125-137.

Ish-Am, G. Eisikowitch, D. 1993. The behaviour of honey bees (Apis mellifera) visiting avocado
(Persea americana) flowers and their contribution to its pollination. Journal of Apicultural
Research 32:175-186.

42
Ish-Am, G. Eisikowitch, D. 1998a. Low attractiveness of avocado (Persea americana Mill.)
flowers to honeybees (Apis mellifera L.) limits fruit set in Israel. Journal of Horticultural
Science & Biotechnology 73:195-204.

Ish-Am, G. Eisikowitch, D. 1998b. Mobility of honey bees (Apidae, Apis mellifera) during
foraging and agricultural pollination. Apidologie 29:209-219.

Ish-Am, G. Eisikowitch, D. 1998c. Quantitative approach to avocado pollination. Proc. Of Third


World Avocado Congress (1995) 46-51.

Ish-Am, G. Gazit, S. 2002. Pollinators of Avocado. Excerpted from: Wysoki, M. Berg, M.A. van
den. Ish-Am, G. Gazit, S. Peña, J.E. Waite G.K.: Pest and Pollinators of Avocado. In:
Peña, J.E. Sharp, J.L. Wysoki M. (eds.): Tropical Fruit Pests and Pollinators. CABI
Publishing, Wallingford UK. 223-293.

Ish-Am, G. Regev, I. Peterman, Y. Lahav, E. Degani, C. Elbatzri, R. Gazit, S. 1999b. Improving


avocado pollination with bumblebees: 3 seasons summary. California Avocado Society
Yearbook 82:119-135.

Ish-Am, G., & Eisikowitch, D. (1993). The behaviour of honey bees (Apis mellifera) visiting
avocado (Persea americana) flowers and their contribution to its pollination. Journal of
Apicultural Research, 32, 175-186. https://doi.org/10.1080/00218839.1993.11101303
Jackson , J. E. 1978. light interception and utilization by orchard systems. Hort Reviews 2:208-
267.
Klein, A.-M.,Vaissiere, B. E., Cane, J. H., Steffan-Dewenter,I., Cunningham, S. A., Kremen, C.,
& Tscharntke, T. (2007). Importance of pollinators in changing landscapes for world
crops. Proceedings of the Royal Society B: Biological Sciences, 274(1608), 303–313.
https:// doi.org/10.1098/rspb.2006.3721
Knight, R.J. Jr 2002. History, distribution and uses. In: Whiley, A.W. Schaffer, B.
Wolstenholme, B.N. (eds.). The Avocado: botany, production and uses. CABI
Publishing, Wallingford UK. 15-37.

43
Lahav, E. Zamet, D. 1999. Flowers, fruitlets and fruit drop in avocado trees. Revista Chapingo
Serie Horticultura 5 Num. Especial. Proc. of Fourth World Avocado Congress (1999) 95-
100.

Lakso, A. (1994). Chapter 2: Apple, in B. Schaffer and P. Andersen (eds.), Handbook of


Environmental Physiology of Fruit Crops, vol. 1, Temperate Crops CRC Press, Florida.
Lesley, J.W. Bringhurst, R.S. 1951. Environmental conditions affecting pollination of Avocados.
California Avocado Society Yearbook 35:169-173.

Lugo, U.R. (2006). Substitución de lardo por grasa vegetal en salchichas: incorporación de pasta
de aguacate. Efecto de la inhibición del oscurecimiento enzimático sobre el color.
Revista de Ciência e Tecnologia de Alimentos, v.26, n.2, p.441-445.
MAC NEISH, R. S. 1964. Ancient Mesoamerican civilization. Science. 143: 531–53
Malerbo-Souza, D.T., Vagner, T., Rodrigues, S., & Sousa, F. (2000). Pollination in avocado
flowers (Persea americana Mill.). Acta Scientiarum - Agronomy, 22(4), 937-941.
McGregor, S.E. 1976. Insect pollination of cultivated crop-plants. U.S.D.A. Agriculture
Handbook No. 496:93-98.

Mickelbart, M.V., Miller, R., Parry, S., Arpaia, M.L. and Heath, R. (2000) Avocado leaf surface
morphology. California Avocado Society Yearbook 84, 139–150.
Middleton, S. (2007). Maximising the light interception and productivity of Australian apple
orchards. Apple and Pear Australia Ltd, Future Orchards notes, viewed 11 November
2018, <https://apal.org.au/wp-content/uploads/2013/07/fo-ow-0207-light-interception-
productivity.pdf>
MIJARES, P., AND L. LÓPEZ. 1998. Variedades de aguacate y su producción en México.
Memoria Fundación Salvador Sánchez Colín CICTAMEX S.C., México.
Monteith, J. (1977) Climate and the efficiency of crop production in Britain. Philosophical
Transactions of the Royal Society B, vol. 281 (980), viewed 16 November 2018,
http://rstb.royalsocietypublishing.org/content/281/980/277
Naveh, E., Werman, M.J., Sabo, E. and Neeman, I. (2002). Defatted avocado pulp reduces body
weight and total heatic fat but increases plasma cholesterol in male rats fed diets with
cholesterol. Journal of Nutrition 132(7) 2015-2018.

44
Palmer, J. (2004). High density planting systems: principles and pitfalls. The Horticulture and
Food Research Institute of New Zealand.

Palmer, J. 1999. computed effects of spacing on light interception and distribution within
hedgerow trees in relation to productivity. acta Horticulturae 114:80-88uction in apple.
hortscience 26:993-999.
Papademetriou, M.K. 1975a. A study of the viability of avocado pollen under natural conditions.
California Avocado Society Yearbook 58:54-56.

Papademetriou, M.K. 1975b. Pollen tube growth in avocados (Persea americana Mill.)
.California Avocado Society Yearbook 58:99-102.

Papademetriou, M.K. 1976. Some aspects of the flower behaviour, pollination and fruit set of
Avocado (Persea americana Mill.) in Trinidad. California Avocado Society Yearbook
60:106-153.

Peña, J.E. (2003). Insectos polinizadores de frutales tropicales: no solo las abejas llevan la miel
al panal. Manejo Integrado de Plagas y Agroecología (Costa Rica), 69, 6-20.
Perez-Balam, J., Quezada-Euan, J. J., Alfaro-Bates, R., Medina, S., McKendrick, L., Soro, A., &
Paxton, R. (2012). The contribution of honey bees, flies and wasps to avocado (Persea
americana) pollination in southern Mexico. Journal of Pollination Ecology, 8, 42–47.
Popenoe, W., 1920. Manual of Tropical and Subtropical Fruits, Macmillan, London, pp 524.
Praloran, J.C. (1970) Le climat des aires d’origine des avocatiers. Fruits 25, 543–557. (in
French.)
Rader, R., Cunningham, S. A., Howlett, B. G., & Inouye, D. W. (2020). Non-bee insects as
visitors and pollinators of crops: Biology, ecology, and management. Annual Review of
Entomology, 65, 391–407. https:// doi.org/10.1146/annur ev-ento- 01101 9-025055
Rodrı´guez-Fragoso, L., Martı´nez-Arismendi, J.L., Orozco-Bustos, D., Reyes-Esparza, J.,
Torres, E., Burchiel, S.W., 2011. Potential risks resulting from fruit/vegetable-drug
interactions: effects on drug-metabolizing enzymes and drug transporters. J. Food Sci. 76,
R112_R124.
Samson, J.A. (1986) Tropical Fruits, 2nd edn. Tropical Agriculture Series. Longman Scientific
and Technical, Harlow, UK, pp. 235–255.
Schaffer , B., and A. W. whiley 2002. environmental phisyology. Pages 135-160 in The
Avocado. CABI.
45
SCHROEDER, C. A. 1968. Prehistoric avocados in California. Cal. Avoc. Soc. 1968 Yrbk. 52:
29–34.
Schroeder, C.A. 1954. Some aspects of pollination in the avocado. California Avocado Society
Yearbook 38:159-162.

Schroeder, C.A. 1955. Pollen production in the avocado. California Avocado Society Yearbook
39:184-186.

SCORA, R. W., AND B. O. BERGH. 1992. Origin of and taxonomic relationships within the
genus Persea. Pages 505–514 in C. J. LOVATT, ED., Proceedings of the Second World
Avocado Congress. Pebble Beach, California
Sedgley, M. (1979). Light-microscope study of pollen-tube growth, fertilization and early
embryo and endosperm development in the avocado varieties Fuerte and Hass. Annals of
Botany, 44, 353-359.
Sedgley, M. 1977. Reduced pollen tube growth and the presence of callose in the pistil of the
male floral stage of the avocado. Scientia Horticulturae 7:27-36.

Sedgley, M. 1980. Anatomical investigation of abscised avocado flowers and fruitlets. Annals of
Botany 46:771-777.

Sedgley, M. Grant, W.J.R. 1983. Effect of low temperatures during flowering on floral cycle and
pollen tube growth in nine Avocado cultivars (Persea americana). Scientia Horticulturae
18:207-213.

Shoval, S. 1987. Pollination rate and pollen tube growth of avocado, in relation to yield. M. Sc.
Thesis, The Hebrew Univ., Rehovot, 161 p. (In Hebrew, English Abstract).

SIAP. Servicio de Información y Estadística Agroalimentaria y Pesquera. 2007. México.


www.siap.gob.mx/aagricola_siap/icultivo/index.js p (acessed 23 September 2006).
Silva, T.A. and Ledesma, N. (2014) Avocado History, Biodiversity and Production. In:
Nandwani, D., Ed., Sustainable Horticultural Systems, Sustainable Development and
Biodiversity, Springer International Publishing, Switzerland, 157-20

Smith, C.E. Jr 1966. Archaeological evidence for selection in avocado. Economic Botay 20:169-
175.

46
Smith, C.E. Jr 1969. Additional notes on pre-conquest avocados in Mexico. Economic Botany
23:135-140.

Stadler, J.D. and Stassen, P.J.C. 1985. Pruning and training deciduous fruit trees: 1. Lighting,
density and pruning procedures. FFTRI Information Bulletin No. 531.

Stassen, P. J. C., B. Snijder , and Z. J. Bard 1999. Results obtained by pruning overcrowded
Avocado orchards. Revista Chapingo Serie Horticultura 5:165-171.
Statistics Division of the Food and Agriculture Organization for the United Nations, 2013.
FAOSTAT data. Available from: ,http://faostat3.fao.org/ home/E.
Sterne, R.E., Kaufmann, M.R. and Zentmyer, G.A. (1977) Environmental effects on transpiration
and leaf water potential in avocado. Physiologia Plantarum 41, 1–6.
STOREY, W. B., B. BERGH, AND G. A. ZENTMYER. 1986. The origin, Indigenous range,
and dissemination of the avocado. Cal. Avoc. Soc. 1986 Yrbk. 70: 127–133.
Storey, W.B., Bergh, B. and Zentmyer, G.A. (1986) The origin, indigenous range and
dissemination of the avocado. California Avocado Society Yearbook 70, 127–133.
Stout, A.B. 1923. A study in cross-pollination of avocado in southern California. California
Avocado Association Annual Report 7:29-45.

Stout, A.B. 1933. The pollination of avocados. Florida Agricultural Experimental Station
Bulletin 257:1-44.

Thorp , T. G., and B. Stowell 2001. Pruning Height and Selective Limb Removal Affect Yield of
Large ‘Hass’Avocado Trees. Hortscience 36(4):699–702.
Tomer, E. Gottriech, M. Gazit, S. 1976. Defective ovules in avocado cultivars. Journal of the
American Society of Horticultural Science 101:620-623. Vithanage, H.I.N.V. 1990. The
role of the European honeybee (Apis mellifera L.) in avocado pollination. Journal of
Horticultural Science 65:81-86.

USDA [United States Dept of Agriculture]. (2010). Nutrient Database. http://www.nal.usda.gov/.


Vithanage, V. (1990). The role of the European honey bee (Apis mellifera L.) in avocado
pollination. Journal of Horticultural Science and Biotechnology, 65(1), 81-86.
Weinberger, K., Lumpkin, T. A. (2007). Diversification into horticulture and poverty reduction:
a research agenda. World Development 2007, 35, 8:1464–1480.

47
Whiley , A. W. 2002. Crop managemant. Pages 231-258 in B. N. wolstenholme editor. The
Avocado: Botany ,Production And Uses.
Whiley, A.W. and Schaffer, B. (1994) Avocado. In: CRC Handbook of Environmental
Physiology of Fruit Crops Vol. II. (Eds) B. Schaffer and P.C. Anderson, CRC Press Inc.,
Boca Raton, Florida, pp. 3-35.

Wolstenholme, B. N. (1987). Theoretical and applied aspects of Avocado yield as affected by


energy budgets and carbon partitioning. South African Avocado Growers' Association
Yearbook 10:58-61.
Wolstenholme, B.N. (1986) Energy costs of fruiting as a yield-limiting factor, with special
reference to avocado. Acta Horticulturae 175, 121–126.
Wolstenholme, B.N. and Whiley, A.W. (1998) Strategies for maximizing avocado productivity –
an overview. In: Proceedings of the Third World Avocado Congress. Tel Aviv, Israel, pp.
61–70.
Wolstenholme, B.N. and Whiley, A.W. (1999) Ecophysiology of the avocado (Persea
americana Mill.) tree as a basis for pre-harvest management. Revista Chapingo Serie
Horticultura Núm. Especial V, 77–88.
Woolf , A. B., I. B. Ferguson , L. C. Requejo-Tapia , and L. Boyd 1999. Impact of Sun exposure
on harvest quality of ‘Hass’ Avocado fruit. Revista Chapingo Serie Horticultura 5:353-
358.
Woyessa G, Berhanu T (2010). Trends of avocado (Persea Americana M.) Production and Its
Constraints in Mana Woreda, Jimma Zone: A Potential Crop for Coffee Diversification.
Trends in Horticultural research, 2010 ISSN 1996-0735 / DOI: 10.3923/thr.2010.
Zekarias S (2010). Avocado Production and Marketing in South Western Ethiopia. Trends Agric.
Econ. 3(4):190-206, 204. ISSN 1994-7933, 2010 Asian Network for Scientific
Information.

48
49

You might also like