ABSTRACT
African pear (Dacryodes edulis (G. Don) HJ Lam) commonly called bush butter is an indigenous multi-purpose but threatened food fruit tree that supports the livelihood of many households in Nigeria. It serves as a food supplement which is normally available during hunger period. Also its medicinal, nutritional and ecological values are highly cherished. . Unfortunately, the species is still mostly collected from the forests and is yet to be developed. This is largely because most of the information necessary for its selection and improvement is presently seriously lacking. Field study trips were therefore conducted in five states of southeastern and south – south Nigeria (Latitude 050 30’N; Longitude 07024’E; Altitude 440m) during the normal flowering, fruiting and ripening period for three (3) consecutive years to evaluate the flowering phenology and effect of application of exogenous growth regulators on African pear. The different types (cultigens) of the species growing in the forests, distant and nearby farms, including homesteads were used. The exogenous growth regulators experiment, the site was D. edulis orchard at IMSU farm. Due to the fact that many tropical fruit trees flower profusely and majority produce many fruits at the onset. Later most of these fruits if not all may be dropped, posing economic problems for the owners. African pear (D. edulis (G. Don) H.J.Lam) is a typical example. The researcher evaluated four growth regulator treatments and calculated their percentage concentration (%). These were Distilled water (control) (T1), 4% Urea (T2), 4%Giberellic acid(T3 ) and 50 % Coconut Water (T4 ). Sixteen mature African pear trees planted seventeen (17) years ago were used for the study and there were four (4) trees in each block. The treatments were applied three (3) weeks before normal flowering time and at two weeks after flowering . Both field and effect of growth regulators characterization studies were made on the variations observed within the species and between the years and location. All were laid out in randomized complete block design. The data were collected and analyzed using analysis of variance and regression/correlation in Stat graphics 16.0 software package. Standard errors and percentages were calculated where appropriate. 450trees and 18,000 fruits were sampled. Parameters collected include time for flowering, fruiting, and fruit set. Important pomological characteristics such as fruit weight, pulp weight, seed weight, fruit length, fruit longitudinal circumference etc. Also collected were floral parameters such as number of inflorescence per flowering branch-let, inflorescence length, number of inflorescence per panicle etc and vegetative parameters like vine length, rachis length, number of leave per ranch etc were also collected.  Based on the observed floral attributes, two distinct AP tree species were identified – Female tree, and Male and hermaphrodite tree. Base on pomological attributes, three distinct types of the species were identified, large (big), medium and small. Ten different ripped fruit colors were identified, four unripe fruit colors, time of flowering, fruiting and fruit set varied within location and between locations. Also significant differences were recorded in fruit weight, length, diameter, longitudinal circumference, number of seed, seed weight, and inflorescence length respectively. Significant (p<0.05) differences were observed in mean seed number at Imo state which ranged from 2.46 -9.14, 7.81- and 8.49 – 11.48 in 2013, 2014 and 2015 respectively. The mean number of fruit per inflorescence were statistical (p<0.05) not similar. This ranges from 0.74- 2.73 in 2013, 1.67-3.08 in 2014. Similarly statistical differences were observed in seed weight, leaf length in Imo location, fruit length, seed weight, inflorescence per panicle in Abia State. Also, time of flowering, fruiting and fruit set varies both within and between locations. In terms of flowering and fruiting habit, AP can be seen as an annual since 57% to 77% of the trees are annual while 23% to 43% are biennial and triennial. Fruits drop were very high- 90% to 95% in all the years and locations.
Application of growth regulators significantly enhanced flowering, fruiting and fruit set. The best fruit retention of 76.56 % was obtained by T4Â which was however, not significantly (p< 0.05) different from 72.75% fruit set obtained using T3. The control (T1) gave the lowest fruit set of 21.41 % which was however significantly (P < 0.05) different from 66.48% obtained by T2. (NS). Application of 4% Giberellin significantly (P < 0.05) enhances flowering, fruiting and fruit set in African pear (D. edulis). But 50% coconut water was more effective in reducing fruit drop.
Therefore, variations that are of genetic and horticultural interest exist in this multipurpose and endangered tree to warrant a planned selection and improvement program and higher dosage of these growth regulators is encouraged.
TABLE OF CONTENTS
Title Pages                                                                                                                                      i Certification                                                                                                                                           ii
Dedication                                                                                                                                    iii
Acknowledgement                                                                                                                        iv
Table of Contents                                                                                                                          vi
List of Tables                                                                                                                               xii
List of Figures                                                                                                                           xviii
List of Plates                                                                                                                             xxii
Abstract                                                                                                                                    xxiii
CHAPTER ONE
1.0      Introduction                                                                                                                        1
CHAPTER TWO
2.1      Botany and origin of African pear (Dacryodes edulis (G. Don) HJ Lam)                      5
2.2      Importance of African pear                                                                                                6
2.3      Phenology of flowering & fruit set                                                                                   8
2.3.1              Temporal patterns in leafing phenology                                                                 12
2.3.1.1           Community Patterns                                                                                             12
2.3.1.2           Herbivory as a Selective Force in Leaf Flushing                                                 13
2.3.1.3           Leaf Fall                                                                                                                 13
2.3.2              Flowering Phenology                                                                                            14
2.3.2.1           Community Pattern in Flowering Phenology                                            14
2.3.2.2           Species Patterns in flowering phenology                                                              15
2.3.2.3           Environmental Cues for Flowering                                                                       16
2.3.2.4           Competition for Pollinators and Interference between Species.                           16
2.3.2.5           Pre-dispersal Seed Predation                                                                                18
2.3.2.6           Inherent Constraints on Flowering                                                                       19
2.3.3              Fruiting Phenology                                                                                               19
2.3.3.1           Selective pressure on timing                                                                                  19 2.3.3.2          Mast Fruiting                                                                                                        20
2.3.3.3            Keystone Fruit Providers                                                                                       21
2.4                  Floral Physiology                                                                                                   21
2.4.1               Vegetative Growth and Floral Initiation                                                              27
2.4.2Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â The Role of Plant Growth Regulators (PGRs) Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 28
2.4.3               The role of Carbohydrates                                                                                    30
2.5.0               Fruit Physiology                                                                                                    31
2.5.1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Regulation of fruit set and growth. Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 32
2.6.0               Exogenous Substances of Flowering and Fruit Set                                               32
CHAPTER THREE
3.0                Materials and Methods                                                                                                        37
3.1                Location and experimental conditions                                                                  37
3.2.0Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â EXPERIMENT 1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 39
3.2                  Flowering Phenology Evaluation and Characterization Studies                         39
3.2.1              Flowering Phenology and Field Survey Studies                                                   39
3.2.1.1            Experimental Materials and Characterization Studies for Field Survey              39
3.2.1.2Â Â Â Â Â Â Â Â Â Â Â Â Data Collection on Flowering Phenology Evaluation
and Characterization Studies for Field Survey                                                 40
3.2.1.2            Statistical Design and Analysis                                                                         43
3.3.0Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â EXPERIMENT 2Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 46
3.3.1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Evaluation of Effects of Exogenous Substances
Application on African pear Floral Initiation. Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 46
3.3.1.1            Experimental Materials and Procedure                                                                46
3.3.1.2            Experimental Design and Treatment                                                      46
3.3.1.3            Treatments and Preparation Application                                                             47
3.3.1.1            Data Collection                                                                                                  47
3.3.1.1            Experimental Design and Analysis                                                                     47
3.4.0Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â EXPERIMENT 3Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 49
3.4.1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Evaluation of Effects of Exogenous Substances
Application on African Pear Fruit Set. Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 49
3.4.1.1            Experimental Materials and Procedures                                                             49
3.3.1.2            Experimental Design and Data Analyses                                                            51
CHAPTER FOUR
4.0                  Results                                                                                                                52
4.1.0Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Experiment 1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 52
4.1.1               Flowering Phenology and Characterization Studies                                       52
4.1.1.1            Phenology of Flowering and Fruiting                                                                  52
4.1.1.2            Flower to Anthesis                                                                                               56
4.1.1.3           Anthesis to Fruit Formation                                                                             59
4.1.1.4            Fruiting Mean Intensity                                                                                        62
4.1.1.4            Fruit Formation to Fruit Set                                                                                  65
4.1.1.5Â Â Â Â Â Â Â Â Â Â Â Â Percentage Mean Number of Fruits Formed(dropped)/ Retained (Matured) Â Â Â Â Â Â 68
4.1.1.7            Ripped Fruit Colors                                                                                           71
4.1.1.8            Unripe Fruit Colors                                                                                          71
4.1.1.9            Fruiting Habit                                                                                                   75
4.1.1.10          Branching Habit                                                                                               77
4.1.1.11          Fruit Shape                                                                                                       79
4.1.1.12          Flushing Phenology                                                                                              81
4.1.1.12.1       Imo state Locations                                                                                          81
4.1.1.12.2       Rivers State                                                                                                          82
4.1.1.12.3       Abia State                                                                                                            82
4.1.1.12.4       Delta State                                                                                                           83
4.1.1.12.5       Enugu State                                                                                                         83
4.1.2.0            Pomological Characterization Studies                                                                89
4.1.2.1            Location: Imo State                                                                                             89
4.1.2.1.1         Sub Locations: (Umuduru Egbeaguru, IMSU (Uratta), and Logara.                    89
4.1.2.1.1.1      Fruit Weight                                                                                                          89
4.1.2.1.1.2      Pulp weight                                                                                                            90
4.1.2.1.1.3      Pulp Thickness                                                                                                      90
4.1.2.1.1.4      Fruit Longitudinal Circumference                                                                        91
4.1.2.1.1.5      Fruit Length                                                                                                          92
4.1.2.2.0         Location: Abia State                                                                                           101
4.1.2.2.1         Sub Location (Oboro, Alayi and Ngwa).                                                          101
4.1.2.2.1.1Â Â Â Â Â Â Fruit Weight:Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 101
4.1.2.2.1.2      Pulp Weight                                                                                                        102
4.1.2.2.1.3      Pulp Thickness                                                                                                     102
4.1.2.2.1.4      Fruit Longitudinal Circumference                                                                      103
4.1.2.2.1.5      Fruit Length                                                                                                        104
4.1.2.2.1.6Â Â Â Â Â Â Â Â Number of seed:Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 105
4.1.2.3.0         Location: Delta state                                                                                         114
4.1.2.3.1Â Â Â Â Â Â Â Â Â Sub Locations: (Anwai, Ugbolu and Ogwashiukwu)Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â 114
4.1.2.3.1.1      Fruit weight                                                                                                    114
4.1.2.3.1.1      Pulp weight                                                                                                       115
4.1.2.3.1.2      Pulp Thickness                                                                                                    115
4.1.2.3.1.3      Fruit Longitudinal Circumference                                                                     116
4.1.2.3.1.4      Fruit Length                                                                                                         117
4.1.2.4.0         Location: Enugu State                                                                                       127
4.1.2.4.1         Sub Locations: Nsukka, Oba, and Amachala.                                                    127
4.1.2.4.1.1      Fruit Weight                                                                                                    127
4.1.2.4.1.2      Pulp weight                                                                                                     127
4.1.2.4.1.3      Pulp Thickness                                                                                                    128
4.1.2.4.1.4      Fruit Longitudinal Circumference                                                                      129
4.1.2.4.1.5      Fruit Length                                                                                                        130
4.1.2.5.0         Location: Rivers state                                                                                        140
4.1.2.5.1         Sub Locations: Etche, Elele, and Obiebe.                                                           140
4.1.2.5.1.1      Pulp weight                                                                                                     140
4.1.2.5.1.2      Pulp Thickness                                                                                                    141
4.1.2.5.1.3      Fruit Longitudinal Circumference                                                                      142
4.1.2.5.1.4      Fruit Length                                                                                                        14
4.1.3.0            Vegetative Morphology                                                                                    154
4.1.4.0            Nature of Inflorescence                                                                                   154
4.2.0Â Â Â Â Â Â Â Â Â Â Â Â Â Â EXPERIMENT 2
4.2.1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Effect of growth regulators on African pear
(Dacryodes edulis (G. Don) HJ Lam) flowering and fruiting.                         156
4.3.0               Experiment 3.                                                                                                 158
4.3.1Â Â Â Â Â Â Â Â Â Â Â Â Â Â Â Effect of Growth Regulators on African pear
(D. edulis (G. Don) HJ Lam) Fruit Set and Fruit Drop                                       158
CHAPTER FIVE
5.0      Discussion                                                                                                                      164
5.1      Flowering Phenology and evaluation Characterization studies                                     164
5.2Â Â Â Â Â Â Evaluation of xogenous Substance Application on Floral Initiation,
Fruiting and Fruit Set Characterization                                                                         169
Conclusion                                                                                                                   171
Recommendation                                                                                                           173
References                                                                                                                       174
Appendix                                                                                                                         188
CHAPTER ONE
INTRODUCTION
The African pear (AP) (Dacryodes edulis (G. Don) HJ Lam) also called bush butter is one of the most important indigenous African Tropical Fruit Trees Species (TFTS). It is a multipurpose tree used as food, medicine, livestock feed and ornamental purposes. The fruit is a source of high quality nutritious, non-greasy natural vegetable oil (44 – 67 %) with possibly numerous industrial uses. The African pear also helps in improving the milk quality of nursing mothers (Okorie et al., 2006). African pear fruit is eaten with maize as butter is eaten with bread. The flowers are useful in apiculture. In recent years there has been a growing awareness of the potentials of African pear as a major source of plant oil; mainly amino acids and triglycerides (Okorie et al., 2006). Despite all these benefits, the AP has not been researched on like other trees because of some problems and constraints such as high flower abortion, propagation constraints, irregular flowering and high post-harvest loses.
In Nigeria, the horticultural fruit trees that have received more research attention are the exotic fruit trees such as citrus (Citrus sinensis cv) and mango (Magnifera indica L). The Few indigenous fruit trees that had been studied include oil palm (Elaeis guineensis Jacq.) and the kola (Cola acuminate (P. Beauv.) Schott & Endl and Cola nitida (Vent) A. Chev.) (Okorie et al., 2000). African pear is little known outside its place of origin (Okorie, 2001).
The long term neglect of TFTS in the area of research, including the African pear (AP) has given rise to a conspicuous lack of information in the horticultural and pomological attributes of the species. This has greatly hampered efforts in their development and optimum utilization. If the environmental and humanitarian dimensions are to be important considerations in future agricultural development agenda, the bridging of the information gap on tropical fruit tree species is a necessity (Okorie et al., 2000). There is a great deal of intra- specific variations in the flowering phenology and fruit set of the AP to warrant the initiation of a selection and improvement programme. But this can only effectively commence when the peculiar abortion problems of the African pear are adequately addressed. This may be achieved through the application of exogenous substances such as auxin, cytokinin, gibberallin, ethylene and abscisic acid.
Knowledge of the AP fruit phenology especially as it relates to flowering and fruit set in the crop is particularly important for the development and improvement of the species.  Phenology is defined as the study of developmental timing in relation to the calendar (Lienth, 1974). This knowledge is of essence because of the following reasons:
- In an environment, it is necessary to match the life cycle of the crop to the length of the growing season for optimal crop yield. Such information is needed to develop better cropping systems so that high and/or stable productivity can be achieved.
- Introduction of improved genotypes or new crops into regions is largely determined by temperature and phenology (Aitken, 1974).
- Phenology is an essential component in whole crop simulation models which can be applied to specific developmental processes to maximize yields.
Application of hormones, particularly to crops plays a key role in increasing yield. These plant hormones also called phytohormones or exogenous substances are chemicals that help in the growth, development and functioning of plants. They also help in the formation of leaves, flowers, stems, and fruits. Most importantly, some of these hormones help in inducing flowering in plants and preventing fruit abortion (Freemen, 1979). Furthermore, these growth regulators (Urea, Coconut water and gibberelin) carry out vital biochemical reactions that are required for plant survival. These hormones help in determining the sex of flowers, colour of fruits and leaves. They help in the formation of tissues, respiration, energy production and even plant longivity and death.
Citrus and mango growers have been able to improve production and yields by manipulating the phenology and physiology of these crops through the application of hormones (Ali & Lovatt, 1994). It has been reported that sweet oranges may develop two hundred and fifty thousand (250,000) flowers per tree in a bloom season, but only a small amount of these flowers become mature fruits (Domingo et al., 2007). Fruit drop in mango is too high, amounting to about 99% at various stages of growth (Ram, 1983). Such research works is lacking in African indigenous tropical fruit trees species especially African pear.
It has been observed that most African pear plants flower profusely during their flowering season, but majority of these flowers are aborted during fruit-set (Okorie, 2001). In the temperate zone, information on the application of exogenous substances to prevent fruit abortion abounds, while in the tropics, there is scarcity of information on flowering phenology and the effects of application of exogenous substances on flowering and fruit set on African indigenous TFTS. Reduction in floral abortion and fruit drop through the application of exogenous substances may result to significant yield increases in African pear.
The growth and development of plants which comprise germination, seedling growth, flowering and fruiting could be influenced positively or negatively by exogenously applied substances. These substances include growth hormones and other secondary metabolites (Erclishi and Turkai, 1998).
Phenology plays a key role in the survival of any plant. For example, in the manifestation of the physiological aspect of any plant, the timing of such event as flowering is very critical.
How and why does Dacryodes edulis (G. Don) HJ Lam (African pear) differ in their flowering phenology?
How does the variations in flowering phenology affect fruit set within the species?
How does the variations in floral morphology and physiology (e.g. relative proportion of male, female and hermaphrodite flowers) or time of flowering affect phenology and fruit set between and within African pear (AP) fruit types?
What factors are responsible for both high percentage floral abortion and prematurity fruit drop observed within the species?
How can we improve on the percentage flowering of African pear while reducing the high incidence of floral abortion and fruit drop observed within species?
Can flowering and fruit set in African pear be improved upon by the application of exogenous substances, and to what extent?
This work was therefore designed to address the above questions.  The timing of life history events may be largely influenced by evolutionary constraints with selection merely modifying a predetermined underlying pattern as with all other genetically controlled characteristics (Chapman et al.,1999).
Therefore, there is the need to study the phenology of flowering in the African pear and the effects of application of exogenous substances on African pear (Dacryodes edulis (G. Don) HJ Lam) flowering and fruit set. The overall objective is not only to provide basic information required for the improvement of the species but also to improve fruit yields within the species.
The specific objectives of this project are as follows:
- To study and evaluate the phenology of flowering in African pear (Dacryodes edulis (G. Don) HJ Lam).
- To evaluate and assess the intraspecific variations in the flowering phenology and fruit set of the African pear fruit types.
- To evaluate the effects of application of exogenous substances (Gibberellin, urea, and coconut water) on flowering and fruit set in African pear.
- To determine the appropriate exogenous substance application level most favorable for increased flowering and fruit set in African pear species.
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