Biofuels From Food Waste: Applications of Saccharification Using Fungal Solid State Fermentation 1st Edition Antoine Prandota Trzcinski

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Biofuels from Food Waste
Applications of Saccharification Using
Fungal Solid State Fermentation
Antoine P. Trzcinski
CRC Press
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Library of Congress Cataloging-in-Publication Data
Names: Trzcinski, Antoine Prandota, author.

Title: Biofuels from food waste : applications of saccharification using
fungal solid state fermentation / Antoine Prandota Trzcinski.
Description: Boca Raton : CRC Press, 2017. | Includes bibliographical references.
Identifiers: LCCN 2017014056 | ISBN 9781138093720 (acid-free paper)
Subjects: LCSH: Biomass energy. | Food waste. | Fungi--Biotechnologoy.
Classification: LCC TP339 .T78 2017 | DDC 662/.88--dc23
LC record available at https://lccn.loc.gov/2017014056
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Contents
Author........................................................................................................................ix
Chapter 1 Bioconversion of Food Wastes to Energy.............................................1

1.1 Introduction................................................................................1
1.2 Ethanol Production.....................................................................5
1.2.1 Pretreatments................................................................. 5
1.2.2 Saccharification............................................................. 5
1.2.3 Process Configurations.................................................. 6
1.2.4 Other Strategies to Improve Ethanol Yield................... 6
1.2.5 Large-Scale Ethanol Production from Food Waste.......9
1.3 Hydrogen Production.................................................................. 9
1.3.1 Substrate Composition..................................................9
1.3.2 Pretreatments............................................................... 13
1.3.3 Process Configurations................................................ 13
1.4 Methane Production................................................................. 15
1.4.1 Single-Stage Anaerobic Digestion.............................. 15
1.4.2 Two-Stage Anaerobic Digestion.................................. 15
1.4.3 Reactor Configurations................................................ 21
1.5 Biodiesel Production................................................................. 22
1.6 Conclusions............................................................................... 22
Chapter 2 Platform Chemical Production from Food Wastes.............................25

2.1 Introduction..............................................................................25
2.2 Platform Chemical Production................................................. 27
2.2.1 Lactic Acid.................................................................. 27
2.2.2 Citric Acid................................................................... 32
2.2.3 Succinic Acid.............................................................. 33
2.2.4 Polyhydroxyalkanoates................................................ 38
2.2.5 Single Cell Oils........................................................... 42
2.2.5.1 Single Cell Oil Production Using
Hydrophilic Feedstock................................. 43
2.2.5.2 Single Cell Oil Production Using
Hydrophobic Feedstock............................... 43
2.3 Other Chemicals.......................................................................46
2.4 Conclusions...............................................................................46
Chapter 3 Enzyme Production from Food Wastes............................................... 49

3.1 Introduction.............................................................................. 49
3.2 Enzyme Production.................................................................. 50
v
vi Contents
3.2.1 Amylases..................................................................... 52
3.2.2 Lignocellulolytic Enzymes.......................................... 54
3.2.3 Pectinolytic Enzymes..................................................60
3.2.4 Proteases...................................................................... 63
3.2.5 Lipases.........................................................................64
3.3 Conclusions...............................................................................66
Chapter 4 Enhanced Glucoamylase Production by Aspergillus awamori

Using Solid State Fermentation........................................................... 69
4.1 Introduction.............................................................................. 69
4.2 Materials and Methods............................................................. 70
4.2.1 Materials...................................................................... 70
4.2.2 Methods....................................................................... 70
4.2.2.1 Effect of Particle Size on Solid State
Fermentation................................................ 70
4.2.2.2 Experimental Design for
Enzyme Production..................................... 71
4.2.2.3 Solid State Fermentation and
Enzyme Extraction...................................... 71
4.2.2.4 Glucoamylase Assay.................................... 71
4.2.2.5 Statistical Analysis...................................... 72
4.2.2.6 Model Validation......................................... 73
4.2.2.7 Analytical Methods..................................... 73
4.3 Results and Discussion............................................................. 73
4.3.1 Effects of Substrate on Glucoamylase Production...... 73
4.3.2 Effects of Temperature................................................ 75
4.3.3 Effects of Particle Size................................................ 77
4.3.4 Effects of Substrate Loading....................................... 77
4.3.5 Volume Reduction of Food Waste after
Enzymatic Hydrolysis................................................. 79
4.3.6 Utilization of Crude Enzymes for Food Waste
Saccharification........................................................... 79
4.3.7 Optimization of Glucoamylase Production
by Response Surface Methodology............................. 81
4.3.8 Validation of the Response Model.............................. 87
4.4 Conclusions............................................................................... 87
Chapter 5 Enhancing the Hydrolysis and Methane Production Potential of

Mixed Food Wastes by an Effective Enzymatic Pretreatment........... 89
5.1 Introduction.............................................................................. 89
5.2 Materials and Methods.............................................................90
5.2.1 Cake Waste for the Production of Fungal Mash.........90
5.2.2 Production of Fungal Mash.........................................90
5.2.3 Hydrolysis of Food Waste........................................... 91
Contents vii
5.2.4 Anaerobic Digestion of Enzymatically

Pretreated Food Waste................................................ 91
5 2.5 Analytical Methods..................................................... 91
5.2.6 Data Analysis.............................................................. 93
5.3 Results and Discussion............................................................. 93
5.3.1 Production of Glucose and Free Amino Nitrogen
from Food Waste Pretreated with Fungal Mash......... 93
5.3.2 Release of Soluble COD in Food Waste
Pretreatment with Fungal Mash..................................96
5.3.3 Anaerobic Digestion of Food Waste Pretreated
with Fungal Mash........................................................96
5.4 Conclusions and Future Prospects............................................99
References.............................................................................................................. 101
Index....................................................................................................................... 121
Author
Antoine P. Trzcinski, PhD, received a PhD from the Chemical Engineering
Department of Imperial College London. Dr. Trzcinski developed a novel process
for producing biogas from municipal solid waste and for the treatment of landfill
leachate. In 2009, he carried out research and development at a pilot scale on the
production of value-added products from algae at Manchester University. He worked
extensively on liquid and solid state fermentation processes using biomass such as
sugarcane bagasse, rapeseed meal, coffee waste, waste glycerol, wheat bran, and
soybean residue for the production of sugars, ethanol, enzymes, and biodiesel using
integrated biorefinery concepts. His research interests include fouling mitigation in
membrane bioreactors, characterization of soluble microbial products, identifica-
tion of bacterial and archaeal strains, pharmaceutical and antibiotic removal from
wastewater, fate of nanoparticles in the environment, and bioelectro stimulation of
microbes to improve bioprocesses through interspecies electron transfer (IET). In
2016, he joined the University of Southern Queensland in Australia as a lecturer and
teaches environmental engineering, environmental engineering practice, hydraulics,
solid and liquid waste treatment, and applied chemistry and microbiology as well as
continuing his research in these fields.
ix
1 Bioconversion of Food
Wastes to Energy
1.1 INTRODUCTION
Food waste (FW) is organic waste discharged from various sources including food
processing plants, and domestic and commercial kitchens, cafeterias, and restau-
rants. According to FAO (2012), nearly 1.3 billion tonnes of foods including fresh
vegetables, fruits, meat, and bakery and dairy products are lost along the food supply
chain. The amount of FW has been projected to increase in the next 25 years due
to economic and population growth, mainly in Asian countries. For example, the
annual amount of urban FW in Asian countries could rise from 278 to 416 million
tonnes from 2005 to 2025 (Melikoglu et al., 2013b). Typical foods wasted in Asia–
Pacific countries and around the world are summarized in Table 1.1 (FAO, 2012).
FW is traditionally incinerated with other combustible municipal wastes for
generation of heat or energy. It should be recognized that FW contains high lev-
els of moisture, which may lead to the production of dioxins during its combustion
together with other wastes of low humidity and high calorific value (Katami et al.,
2004). In addition, incineration of FW can potentially cause air pollution and loss
of chemical values of FW. These suggest that an appropriate management of FW is
strongly needed (Ma et al., 2009a). FW is mainly composed of carbohydrate poly-
mers (starch, cellulose, and hemicelluloses), lignin, proteins, lipids, organic acids,
and a remaining, smaller inorganic part (Table 1.2). Hydrolysis of carbohydrate in
FW may result in the breakage of glycoside bonds with releasing polysaccharides as
oligosaccharides and monosaccharides, which are more amenable to fermentation.
Total sugar and protein contents in FW are in the range of 35.5%–69% and 3.9%–
21.9%, respectively. As such, FW has been used as the sole microbial feedstock for
the development of various kinds of value-added bioproducts, including methane,
hydrogen, ethanol, enzymes, organic acid, biopolymers, and bioplastics (Han and
Shin, 2004; Rao and Singh, 2004; Wang et al., 2005a; Sakai et al., 2006; Yang et al.,
2006; Ohkouchi and Inoue, 2007; Pan et al., 2008; Koike et al., 2009; He et al.,
2012b; Zhang et al., 2013b). Fuel applications ($200–400/ton biomass) usually cre-
ate more value compared to generating electricity ($60–150/ton biomass) and animal
feed ($70–200/ton biomass). Due to inherent chemical complexity, FW also can be
utilized for production of high-value materials, such as organic acids, biodegradable
plastics, and enzymes ($1000/ton biomass) (Sanders et al., 2007). However, it should
be noted that the market demand for such chemicals is much smaller than that for
biofuels (Tuck et al., 2012). Therefore, this chapter intends to review the FW valo-
rization techniques that have been developed for the production of various biofuels,
such as ethanol, hydrogen, methane, and biodiesel.
1
2
TABLE 1.1
Typical Wasted Foods in Several Asia–Pacific Countries and around the Globe
Southeastern New North South
Waste (KT) World Asia Asia Australia Cambodia China Indonesia Japan Malaysia Zealand Korea Philippines Korea Thailand Vietnam
Cereal 95,245 52,374 12,599 1,380 506.1 18,990 4.588 413.4 183.4 28.6 253 215.7 628.4 1,999 2,706
Rice 26,738 22,668 10,792 0.4 506.0 6,046 3.307 139.4 50.2 ND ND 162.7 458.2 1,997 2,478
Sugar 459.9 188.9 151.7 93.6 ND 0.4 ND 20.8 ND ND ND ND ND 151.7 ND
Pulses 2,735 1,134 241.6 36.0 0.9 142.3 38.0 7.1 ND 1.2 10.3 ND 2.0 7.0 8.6
Oil crops 18,424 13,590 2,515 3.9 3.8 9,017 2,238 69.6 1.4 0.1 15.2 ND 12.7 159.4 30.5
Vegetable 616.1 269.3 116.9 ND ND 133.4 ND 13.0 116.9 ND ND ND ND ND ND
oil
Vegetables 81,441 59,949 2,710 54.1 46,9 39,286 755.0 1,224 64.8 73.2 414.2 242.5 1,555 339.5 777.2
Beans 1,049 447.3 218.1 1.1 0.9 49.1 37.2 6.5 ND 0.2 10.3 2.2 1.6 3.7 5.2
Onions 5,891 3,877 186.0 14.6 ND 2,107 99.9 68.1 ND ND 3.5 6.9 139.5 5.5 22.7
Peas 412.7 145.1 2.1 7.2 ND 39.9 ND 0.4 ND 1.1 ND 0.3 0.1 0.1 ND
Tomatoes 12,874 7,415 104.2 ND ND 3,181 85.3 100.7 1.6 9.5 8.3 9.9 57.6 7.3 ND
Potatoes 62,229 12,912 466.1 23.6 ND 7,501 250.0 177 ND 10.9 156.0 34.4 95.3 9.0 83.3
Fruits 53,796 28,328 4,529 30.9 30.5 8,323 2,706 749 89.1 43.4 153.5 1,183 276.6 786.4 531.0
Apples 5,742 4,116 13.2 5.9 ND 3,192 3.1 84.6 ND 22.4 72.8 3.8 49.0 1.2 5.1
Bananas 13,532 8,544 1,896 5.4 7.8 949.3 637.4 213.0 56.1 7.6 ND 901.3 ND 153.7 137
Coconuts 3,038 2,488 2,159 ND ND 20.5 2,066 ND 1.3 ND ND 7.8 ND 69.1 0.9
Pineapples 1,829 579 431.9 ND 2.2 97.7 ND 15.4 ND 0.3 ND 109.9 2.8 189.5 50
Coffee 105.0 33.3 28.3 ND ND 0.033 20.9 ND 0.6 ND ND 6.4 ND ND ND
Milk 16,560 10,887 183.3 ND 1.6 1,447 45 ND 3.8 164.8 4.9 ND 42.4 25.2 9.5
(Continued)
TABLE 1.1 (Continued)
Typical Wasted Foods in Several Asia–Pacific Countries and around the Globe
Southeastern New North South
Waste (KT) World Asia Asia Australia Cambodia China Indonesia Japan Malaysia Zealand Korea Philippines Korea Thailand Vietnam
Cream 33.9 0.1 ND ND ND 0.1 ND ND ND ND ND ND ND ND ND
Butter 84.0 1.7 ND ND ND ND ND ND ND ND ND ND ND 23.1 ND
Animal fats 174.1 1.8 ND ND ND 0.1 ND ND ND ND ND ND ND ND ND
Meat 1,184 183.2 ND ND ND ND ND 107.2 ND ND ND ND 107.2 23.1 ND
Offal 63.0 19.6 ND 8.7 ND ND ND ND ND ND ND ND ND ND ND
Poultry meat 97.5 61.2 ND ND ND ND ND 34.5 ND ND ND ND ND 23.1 ND
Annual waste 0.184 ND 0.130 0.277 0.173 0.061 0.130 0.129 0.113 0.280 0.211 0.130 0.098 0.130 0.130
production
per capita
Bioconversion of Food Wastes to Energy
(T)
Population 7,067 4,175 610 22.9 14.5 1,354 237.6 127.5 29.6 4.5 24.6 92.3 50.0 65.9 88.8
(millions)
Total FW 1300a 278b ≥79.3a ≥6.34a 2.50c 82.80d ≥30.90a 16.40d 3.36a,e ≥1.25a 5.19d ≥12.00a 4.91d ≥8.6a ≥11.55a
(MT)
Source: Reprinted from Fuel, 134, Uçkun, K. E. et al., Bioconversion of food waste to energy: A review, 389–399, Copyright 2014c, with permission from Elsevier.
Note: FW: food waste, T: ton, KT: kilotons, MT: million tonnes.
a Gustafsson et al. (2011)
b Melikoglu et al. (2013b)
c Seng (2010)
d OECD (2007)
e Noor et al. (2013)

3
4
TABLE 1.2
Composition of Mixed Food Waste
Moisture Total Solid Volatile Solid Total Sugar Starch Cellulose Lipid Protein Ash References
79.5 20.5 95.0 NA NA NA NA 21.9 NA Han and Shin (2004)
84.1 15.9 15.2 NA NA NA NA NA NA Kim et al. (2004)
80.0 20.0 93.6 NA NA NA NA NA 1.3 Kwon and Lee (2004)
85.0 15.0 88.5 NA NA 15.5 8.5 6.9 11.5 Rao and Singh (2004)
79.1 20.9 93.2 NA NA NA NA NA NA Ramos et al. (2012)
75.9 24.1 NA 42.3 29.3 NA NA 3.9 1.3 Ohkouchi and Inoue (2006)
87.1 12.9 89.5 NA NA NA NA NA NA Kim et al. (2008c)
80.8 19.2 92.7 NA 15.6 NA NA NA NA Pan et al. (2008)
80.3 19.7 95.4 59.8 NA 1.6 15.7 21.8 1.9 Tang et al. (2008)
82.8 17.2 89.1 62.7 46.1 2.3 18.1 15.6 NA Wang et al. (2008a)
75.2 24.8 NA 50.2 46.1 NA 18.1 15.6 2.3 Wang et al. (2008b)
85.7 14.3 98.2 42.3 28.3 NA NA 17.8 NA Zhang et al. (2008)
82.8 17.2 85.0 62.7 46.1 2.3 18.1 15.6 NA Ma et al. (2009a)
61.3 38.7 NA 69.0 NA NA 6.4 4.4 1.2 Uncu and Cekmecelioglu (2011)
4.4 35.6 NA NA NA NA 8.8 4.5 1.8 Cekmecelioglu and Uncu (2013)
81.7 18.3 87.5 35.5 NA NA 24.1 14.4 NA He et al. (2012a)
81.5 18.5 94.1 55.0 24.0 16.9 14.0 16.9 5.9 Vavouraki et al. (2012)
81.9 14.3 98.2 48.3 42.3 NA NA 17.8 NA Zhang and Jahng (2012)
Source: Reprinted from Fuel, 134, Uçkun, K. E. et al., Bioconversion of food waste to energy: A review, 389–399, Copyright 2014c, with permission
from Elsevier.
Total solid, total sugar, starch, cellulose, lipid, protein, and ash contents are given in wt% on the basis of dry weight. Volatile solid contents are given as
the %VS ratio on total solid basis.
Bioconversion of Food Wastes to Energy 5
1.2 ETHANOL PRODUCTION

Recently, global demand for ethanol has increased due to its wide industrial appli-
cations. Ethanol is mainly used as a chemical feedstock to produce ethylene with a
market demand of more than 140 million tonnes per year, a key material for further
production of polyethylene and other plastics. As such, bioethanol produced from
cheap feedstocks has gained interest (Lundgren and Hjertberg, 2010; International
Renewable Energy Agency, 2013). Traditionally, bioethanol is produced from cellu-
lose and starch rich crops, for example, potato, rice, and sugar cane (Thomsen et al.,
2003). Starch can be easily converted to glucose by commercial enzymes and sub-
sequently fermented to ethanol particularly by Saccharomyces cerevisiae. However,
the hydrolysis of cellulose is more difficult. FW hydrolysis becomes much harder
if large quantities of cellulosic feedstocks are present in FW. Use of abundant and
cheap wastes such as lignocellulosic, municipal, and FW has been explored as alter-
native substrates for ethanol production (Kim and Dale, 2004; Jensen et al., 2011).
1.2.1 Pretreatments
Harsh pretreatment may not be necessary during the conversion of FW to ethanol
prior to enzymatic hydrolysis (Kumar et al., 1998; Tang et al., 2008). Instead, auto-
clave of FW before fermentation is often required for improving product yield and
purity, but at the cost of energy and water consumption. It should be noted that ther-
mal treatment may lead to partial degradation of sugars and other nutritional compo-
nents, as well as side reactions (e.g., Maillard reactions) through which the amounts
of useful sugars and amino acids are reduced (Sakai and Ezaki, 2006). Moreover,
fresh and wet FW appear to be more effective than rewetted dried FW (Kim et al.,
2005). This is mainly due to the decreased specific surface area of the dried sub-
strate, resulting in a decrease in the reaction efficiency between the enzymes and
substrate. Therefore, the utilization of FW without a drying pretreatment is preferred
as long as microbial contamination is manageable. Without thermal sterilization,
acidic condition is needed to prevent microbial contamination and putrefaction (Ye
et al., 2008a; Koike et al., 2009). As such, acid-tolerant ethanol-producing micro
organisms such as Zymomonas mobilis have been employed for the fermentation of
FW (Tao et al., 2005; Wang et al., 2008a).
1.2.2 Saccharification
The conversion efficiency of FW to ethanol depends on the extent of carbohydrate
saccharification as yeast cells cannot ferment starch or cellulose directly into bioeth-
anol (Tubb, 1986). A mixture of α-amylase, β-amylase, and glucoamylase of various
origins is more effective for substrates with higher molecular weight. Pullulanase has
also been added to the list of saccharifying enzymes recently (Tomasik and Horton,
2012). As a direct endo-acting debranching enzyme, pullulanase can specifically
catalyze the hydrolysis of α-1,6-glucosidic linkages of branched polysaccharides
(e.g., pullulan, dextrin, amylopectin, and related polymers), resulting in the release of
linear oligosaccharides. Small fermentable sugars (e.g., maltose, amylose, glucose,
6 Biofuels from Food Waste
maltose syrups, and fructose) can be produced in saccharification process, whereas

cellulases and xylanases including endoglucanase, exoglucanase, β-glucosidase, and
β-xylosidase, can also be employed to improve the hydrolysis of cereals for conver-
sion of starches to glucose (Ducroo, 1987).
Table 1.3 shows the glucose and ethanol yields of different types of FW. The
highest glucose concentration of about 65 g reducing sugar (RS)/100 g FW was
obtained with α-amylase at a dose of 120 U/g dry substrate, glucoamylase (120 U/g
dry substrate), cellulase (8 FPU/g dry substrate), and β-glucosidase (50 U/g dry sub-
strate) (Cekmecelioglu and Uncu, 2013). In a study by Hong and Yoon (2011), a mix-
ture of commercial enzymes consisting of α-amylase, glucoamylase, and protease
resulted in 60 g RS/100 g FW.
1.2.3 Process Configurations

High glucose yield is achievable by increasing enzyme concentration and temperature
at different solid loads, agitation speeds, and hydrolysis times in the saccharifica-
tion processes (Sharma et al., 2007; Ado et al., 2009; Shen et al., 2009; Zhang et al.,
2010). High glucose concentration may result in catabolite repression of the enzymes
(Oberoi et al., 2011b). Therefore, fed-batch and simultaneous saccharification and fer-
mentation methods have been developed for achieving high ethanol yield from FW
(Ma et al., 2009b; Oberoi et al., 2011b). The fed-batch culture has been commonly
employed for the production of high concentration reducing sugars which can be fur-
ther fermented to ethanol (Ballesteros et al., 2002). Compared to batch culture, Yan
et al. (2012b) found that saccharification and subsequent ethanol fermentation were
both improved significantly using fed-batch configuration, for example, the glucose
bioconversion yield reached 92% of its theoretical value. Alternatively, Ssf can be
deployed to mitigate risk of catabolite repression. This combines enzymatic hydroly-
sis and ethanol fermentation into a single operation for keeping the concentration of
enzymatically produced glucose at a low level so as to mitigate inhibition to enzymatic
hydrolysis (Hari Krishna et al., 2001). This combined process can be performed in a
single tank, with lower energy consumption, higher ethanol productivity, in a shorter
processing time using less enzyme (Ballesteros et al., 2002). Optimization of fermen-
tation conditions is vital for the success of the Ssf process as enzymes and fermenting
microorganisms may have different optimum pH and temperatures. In a study by
Hong and Yoon (2011), about 60 g RS and 36 g ethanol were produced from 100 g of
FW in 48-h fermentation. Koike et al. (2009) also reported production of ethanol from
non-diluted FW (garbage) in a continuous Ssf process with an ethanol productivity
of 17.7 g/L h. Ma et al. (2009a) investigated the Ssf process using kitchen garbage
by acid tolerant Zymomonas mobilis without any sterilization. Approximately 15.4 g
sugar per 100 g of garbage and 0.49 g ethanol per grams sugar was obtained within
14 h, giving an ethanol yield of 10.08 g/L h.
1.2.4 Other Strategies to Improve Ethanol Yield

To improve ethanol productivity, various strategies have been explored, includ-
ing use of strains with high ethanol tolerance (He et al., 2009; Wang et al., 2012)
TABLE 1.3
Ethanol Production from Food Wastes
Y Y Y
Duration (g RS/100 g (g/g (g/g P
Waste Method Vessel Type Pretreatment Microorganism (h) FW) FW) RS) (g/h) References
Bakery waste Simultaneous 14 L fermenter None S. cerevisiae 14 54 0.25 0.46 NR Kumar et al.
(1998)
KW Repeated batch 1 L fermenter with None S. cerevisiae 264 12.3 0.06 0.5 3.7 Ma et al. (2007)
Simultaneous 0.8 L working vol. ATCC26602
Mandarin Simultaneous 500 mL flask Drying, steam S. cerevisiae Anr, 24 25.2 0.11 0.4 NR Sharma et al.
waste, explosion Pachysolen (2007)
banana peel tannophilus
FW Separate 500 mL flask 100 mL None S. cerevisiae 16 23.4 0.12 0.49 NR Kim et al.
working vol. KA4 (2008c)

FW Simultaneous Flask with 100 g FW None S. cerevisiae 48 11.25 0.08 NR NR Ma et al. (2008)
KW Separate Tower shaped LAB spraying S. cerevisiae 15 11.7 0.03 0.26 24 Tang et al.
Continuous reactor, 0.45 L strain KF-7 (2008)
working vol.
KW Simultaneous Flask with 100 g FW None S. cerevisiae 67.6 34.8 0.23 NR NR Wang et al.
(2008c)
FW Continuous Fermenter with LAB spraying S. cerevisiae 25 36.4 0.09 0.24 17.7 Koike et al.
Simultaneous 4.3 kg FW KF7 (2009)
FW Simultaneous 1 L fermenter with None S. cerevisiae 48 8.9 0.06 NR 10.08 Ma et al.
0.8 L working vol. KRM-1 (2009a)
KW Repeated batch 250 mL flask 150 mL None Zymomonas 14 15.4 0.07 0.49 10.08 Ma et al. (2008)
Simultaneous working vol. mobilis GZNS1
FW Simultaneous 250 mL flask 200 mL None S. cerevisiae 48 60 0.36 0.22 NR Hong and Yoon
working vol. (2011)
7
(Continued)
8

Ethanol Production from Food Wastes
Y Y Y
Duration (g RS/100 g (g/g (g/g P
Waste Method Vessel Type Pretreatment Microorganism (h) FW) FW) RS) (g/h) References
FW Separate 5 L fermenter with None S. cerevisiae 24 27 0.16 NR 1.18 Kim et al.
working volume of (2011c)
3L
FW Synchronous Fermenter with 200 g None Saccharomyces 352 12.5 NR NR 2.24 Li et al. (2011)
Saccharification FW italicus KJ
Mandarin Simultaneous 100 mL baffled Drying S. cerevisiae 15 52 0.34 NR 3.5 Oberoi et al.
waste flasks (2011a)
Banana peels Simultaneous 100 mL baffled Drying S. cerevisiae 15 37.1 0.32 0.43 2.3 Oberoi et al.
flasks (2011a)
KW Separate 250 mL flask 100 mL None S. cerevisiae 96 50 0.2 0.39 NR Uncu and
working vol. Cekmecelioglu
(2011)
KW Separate 250 mL flask 100 mL None S. cerevisiae 48 64.8 0.23 0.36 NR Cekmecelioglu
working vol. and Uncu
(2013)
Bread waste Separate 300 mL flask 80 g Drying S. cerevisiae 72 37 0.27 NR NR Kawa-Rygielska
bread waste Ethanol Red et al. (2012)
KW Separate (fb) 500 mL flask 200 g None S. cerevisiae 48 29 0.14 0.47 NR Yan et al.
FW H058 (2012a)
Note: NR: not reported, FW: food waste, KW: kitchen waste, RS: reducing sugar, Y: yield, P: productivity, Simultaneous: simultaneous saccharification fermentation,
Separate: separate saccharification fermentation, fb: fed-batch.
and cell recycle through sedimentation or membrane retention (He et al., 2012a).
Recombination of bioethanol-producing strains with the amylase-producing gene
or the development of new strains with improved ethanol tolerance has also been
reported (Li et al., 2011). However, stability of the recombinant gene has not yet been
proven. Cell recycling has been known to significantly improve the performance of
the continuous fermentation process (Wang and Lin, 2010).
1.2.5 Large-Scale Ethanol Production from Food Waste

Pilot and full scale plants for ethanol production from various wastes have been
reported. The pilot study by Kumamoto University and Hitachi Zosen Company
showed that 60 L of ethanol could be produced from 1 ton of municipal solid wastes,
while the residual by-products could be further used for biogas production (Japan-
for-Sustainability, 2013). In Finland, ST1 Biofuel built a network of seven ethanol
plants converting various kinds of wastes to ethanol with a total annual capacity
of 11 ML (Energy Enviro Finland, 2013; ST1, 2013). In Spain, citrus wastes have
been converted to ethanol with a yield of 235 L/ton dry orange peel (BEST, 2013;
Citrotechno, 2013). E-fuel developed a home ethanol system supported with micro-
sensors to convert sugar/starch rich liquid wastes into ethanol for homeowners and
small businesses (E-fuel, 2009). A theoretical estimate based on the data presented
in Tables 1.1 and 1.3 suggests that 36.2, 126.8, and 593 TL (teraliters) of ethanol
might eventually be produced annually in Southeast Asia, Asia, and the world,
respectively.
1.3 HYDROGEN PRODUCTION

Hydrogen (H2) is used as compressed gas and has a high energy yield (142.35 kJ/g).
FW rich in carbohydrates is suitable for H2 production. Table 1.4 summarizes the
recent studies on H2 production from FW. It can be seen that the hydrogen yields
ranged from 0.9 mol H2/mol hexose to 8.35 mol H2/mol hexose (Patel et al., 2012).
The factors such as the composition of FW, pretreatments, and process configura-
tions may affect H2 production.
1.3.1 Substrate Composition
Hydrogen production potential of carbohydrate-based waste was reported to be 20
times higher than that of fat-based and protein-based waste (Show et al., 2012). This
was partially attributed to the consumption of hydrogen toward ammonium using
nitrogen generated from protein biodegradation. Kim et al. (2010) reported that the
H2 yield was maintained at around 0.5 mol H2/mol hexose at the C/N ratio lower
than 20, while the H2 yield was found to drop at higher C/N ratio because of the
increased production of lactate, propionate, and valerate. The H2 yield was signifi-
cantly enhanced and reached to 0.9 mol H2/mol hexose when the C/N ratio was bal-
anced with an alkaline shock.
10
TABLE 1.4
Hydrogen Production from Food Waste
Y
OLR (mol/ Y P
Duration HRT (kg VS/ OLR (kg mol (mL/g (g H2/
Waste Vessel Type Pretreatment Microorganism (day) (day) m3d) COD/m3) Hexose) VS) Lh) References
FW Leaching bed None HSSS 7 5 NR NR NR 160 NR Han and Shin
reactor with 3.8 L (2004)
working vol.
FW with 415 mL bottle with None HSSS 3 Batch NA NA 0.9 67 9.9 Kim et al. (2004)
sludge 200 mL working
vol.
FW 715 mL bottle with None Acidogenic 6 Batch NA NA 1.8 92 6.8 Shin et al. (2004)
500 mL working culture from
vol. CSTR
FW Bioreactor with 3 L None Anaerobic SS 5 NR 8 NR 2.2 125 3.8 Shin and Youn
working vol. (2005)
FW Bioreactor with 3 L None Anaerobic SS 60 5 3 NR 2.4 NR NR Youn and Shin
working vol. (2005)
FW CSTR with 10 L None SS 150 1.3 38.4 64.4 NR 283 19.9 Chu et al. (2008)
working vol.
FW 1 L bioreactor with None Anaerobic SS 2 Batch NA NA NR 57 NR Pan et al. (2008)
500 mL working
vol.
FW 7.5 L bioreactor Heat pretreatment SS 3 Batch NA NA 2.05 153.5 19.2 Kim et al.
with 3 L working (90°C 20 min) (2008c)
vol.
(Continued)
Y
OLR (mol/ Y P
FW ASBR with 4.5 L None HSSS NR SRT: 5.25 NR NR 1.12 80.9 10.2 Kim et al.
working vol. HRT: (2008a)
1.25
KW Bioreactor with 1 L None SS 2 Batch NA NA NR NR 1.0 Lee et al. (2008)
working vol.
FW Rotating drum with None None 30 4 22.65 NR NR 65 NR Wang and Zhao
200 L working (2009)
vol.
Apple 150 mL bioreactor Enzymatic HSSS 2 Batch NA NA NR 134 NR Wang et al.
pomace with 100 mL pretreatment (2010a)
working vol.
FW CSTR 500 L Heat pretreatment HSSS 90 21 NR 12.3–71.3 1.82 NR NR Lee et al. (2010a)
working vol. (100°C 30 min)
KW CSTR with 20 L None SS 59 4 NR NR NR NR 7.1 Lee et al. (2010b)
working vol.
FW SCR with 10 L None HSSS 96 1.9 NR 39 2.5 114 41.3 Lee et al. (2010b)
working vol.
FW ASBR with Alkaline HSSS 200 36 NR NR 0.9 NR NR Kim et al. (2010)
0.15 m3 working pretreatment
vol. (pH 12.5, 1 d)
(Continued)
11
12

Y
OLR (mol/ Y P
FW Bottle with 200 mL Ultrasonication None 14.6 Batch NA NA NR 118 NR Elbeshbishy et al.
working vol. with acid (2011)
FW Bottle with 200 mL None None 3 Batch NA NA 1.79 NR 33.0 Kim et al.
working vol. (2011a)
FW 500 mL bioreactor None HSSS 1 Batch NA NA NR NR 6.6 Mohd Yasin et al.
with 200 mL (2011)
working vol.
FW 300 mL bioreactor None HSSS 2 Batch NA NA NR NR NR Ramos et al.
with 150 mL (2012)
working vol.
FW Bioreactor with Lactate Irradiated R. 1 Batch NA NA 8.35 NR NR Kim and Kim
150 mL working fermentation sphaeroides (2013)
vol. KD131
Note: FW: food waste, KW: kitchen waste, Y: yield, P: productivity, ASBR: anaerobic sequencing batch reactor, SBR: sequencing batch reactor, SS: seed sludge, HSSS:
heat shocked seed sludge, NR: not reported, NA: not applicable.
1.3.2 Pretreatments
Typically, mixed cultures have been employed for H2 production from waste materi-
als. However, hydrogen generated by Clostridium and Enterobacter is often readily
consumed by hydrogenotrophic bacteria (Li and Fang, 2007). Seed biomass is gener-
ally pretreated with heat to suppress hydrogen consumers (Elbeshbishy et al., 2011).
FW itself can be a source of H2-producing microflora. Kim et al. (2008a) have applied
several pretreatments to select microflora for hydrogen production. Lactic acid bacte-
ria are the most abundant species in untreated FW, while H2-producing bacteria are
dominant in the pretreated FW. Heat treatment is effective for suppressing lactate
production and increasing H2/butyrate production. However, heat treatment is likely
to increase costs in large-scale operations. Luo et al. (2010) investigated different
pretreatment methods of inoculums, and concluded that pretreatment would only
have short-term effects on hydrogen production, and the pretreatment is not very
crucial (Wang and Zhao, 2009).
1.3.3 Process Configurations

Various fermentation systems, such as the batch, semi-continuous, continuous, one
or multiple stages, have been developed for production of H2 from FW (Hallenbeck
and Ghosh, 2009). High H2 production rates have been reported in the anaerobic
sequencing batch (ASBR) and up-flow anaerobic sludge blanket (UASB) reactors due
to their high reactor biomass concentrations (Kim et al., 2008a). In these processes,
the solid retention time (SRT) determines the substrate uptake efficiency, microbial
size and composition, and metabolic pathway. A long SRT favors the growth of H2
consumers, while a short SRT may reduce substrate uptake efficiency, active biomass
retention, and subsequently the overall process efficiency. If the optimal SRT could
be achieved at a low hydraulic retention time (HRT), it would enhance the produc-
tivity and technical feasibility of the H2 production process (Wang and Zhao, 2009).
Kim et al. (2008a) investigated the effects of SRT in the range of 24–160 h and
HRT of 24–42 h on hydrogen production from FW. It was found that the maximum
H2 yield of 80.9 mLH2/g volatile solid (VS), equivalent to 1.12 mol H2/mol hexose,
was obtained at SRT of 126 h and HRT of 33 h. Wang and Zhao (2009) obtained a
hydrogen yield of 65 mL H2/g VS at a long SRT of 160 d in a two-stage process. It is
still debatable as for the effect of the organic loading rate (OLR) on bioconversion of
FW to H2. In some studies, lower H2 yields were observed at higher OLRs, whereas
the opposite trend was also reported in the literature. It appears that an optimal OLR
would exist for the maximum H2 yield (Wang and Zhao, 2009). Wang and Zhao
(2009) reported that the hydrogen fermentation pathway became dominant and H2
yield was steady at lower OLR (≤22.65 kg VS/m3 d), while a decrease in the hydro-
lysis rate of substrate and an increase of propionic and lactic acids were observed.
These suggest the possibility of co-production of organic acids if the cost related
to separation is comparable with the value of the products. The inhibitory effect of
organic acids produced at high OLR was also reported by Shin and Youn (2005).
Therefore, it is important to determine the optimum OLR and SRT for improv-
ing H2 production. Acidity of the fermentation medium is another crucial parameter
14 Biofuels from Food Waste
influencing the fermentation efficiency. It had been reported that the optimum pH
for H2 production from organic waste ranged from 4.5 to 6.5 (Kyazze et al., 2007).
The accumulation of fermentation products, that is, CO2, increases the acidity and
then inhibits the microbial growth. Such fermentation products can be removed from
the fermentation medium by simple gas sparging and mixing. The addition of alka-
line or inoculum recycling are also frequently used for pH control (Li and Fang,
2007; Lee et al., 2008; Wang and Zhao, 2009; Kim et al., 2010; Lee et al., 2010a,b).
Compared to the addition of alkali, sludge recirculation is an economically pref-
erable approach for pH control. The long-term stability of a continuous two-stage
process was maintained by recirculating high-alkalinity sludge, for example, at a
OLR of 39 g COD/L d and HRT of 1.9 d, the system was stabilized at 2.5 mol H2/
mole hexose, 114 mL H2/g VS, and 462.5 mL H2/L h over a period of 96 days (Lee
and Chung, 2010).
The bioconversion yield of FW to H 2 production is low, for example, only about
33% of COD in organic materials can be harvested as H 2, while most of the energy
content in the feedstock mainly end up as organic acids, such as acetic, lactic, and
butyric acids. In other words, actual H 2 yield is much smaller than its theoretical
value of 12 mol H 2/mol glucose (Kim and Kim, 2013). As a result, the commercial
value of organic acids—particularly lactic acid—should be further explored. To
improve economic viability of the bioconversion process, H 2 production should
also be combined with the methane, organic acids, and ethanol production pro-
cesses (Lin et al., 2013). Kyazze et al. (2007) reported that the efficiency of the
H 2 production process was improved using the two-stage H 2-methane production
process. Lee et al. (2010a) reported the feasibility of continuous H 2 and CH4 fer-
mentation in a two-stage process using sludge recirculation from the sludge stor-
age tank (denitrification + digestion sludge storage) in a full-scale system. Even
so, only 2.5 mol H 2/mol hexose was obtained due to the limitations of anaerobic
metabolism.
Alternatively, photo-fermentation has also been explored for the conversion of
organic acids to H2. In order to increase the overall H2 yield, a combined dark- and
photo-fermentation system has been proposed. In this process, lactic acid produced
from FW is utilized by photo-fermentative bacteria, particularly purple non-sulfur
bacteria and finally converted to H2 while the remaining residue is converted to CH4
(Show et al., 2012). Overall, via the three-stage fermentation system, 41% and 37%
of the energy content in the FW could be harvested as H2 and CH4, respectively, cor-
responding to the electrical energy yield of 1146 MJ/ton FW (Kim and Kim, 2013).
Lee and Chung (2010) conducted a cost analysis of hydrogen production from FW
using two-phase hydrogen/methane fermentation, and suggested that the abundance
and low cost of FW makes it economically more feasible than the other sources for
H2 production. However, the economic feasibility of process applications from FW
is dependent on the cost of FW collection. Besides, hydrogen production processes
should be combined with an ancillary process, such as methane fermentation, to
achieve complete treatment and disposal of FW. Last, it should also be recognized
that the technological and economic challenges associated with the fermentative H2
production and its purification, storage, and distribution may also slow down the
wide application of bio H2 as green energy.
1.4 METHANE PRODUCTION

The production of biogas, particularly methane via anaerobic processes, is an accept-
able solution for waste management because of its low cost, low production of residual
waste, and its utilization as a renewable energy source (Morita and Sasaki, 2012; Nasir
et al., 2012). In addition to biogas, a produced nutrient-rich digestate can also be used
as fertilizer or soil conditioner. Table 1.5 summarizes the studies pertaining to anaero-
bic digestion of various kinds of FW. Mtz. Viturtia et al. (1989) investigated two-stage
anaerobic digestion of fruit and vegetable wastes, in which 95.1% volatile solids (VS)
conversion with a methane yield of 530 mL/g VS was achieved. In a study by Lee
et al. (1999), FW was converted to methane using a 5-L continuous digester fed with
an OLR of 7.9 kg VS/m3 d, resulting in a 70% VS conversion with a methane yield
of 440 mL/g VS. Gunaseelan (2004) has reported the methane production capacities
of about 54 different fruit and vegetable wastes ranging from 180 to 732 mL/g VS
depending on the origin of wastes.
Feedstock characteristics and process configuration are the main factors affect-
ing the performance of anaerobic digestion (Molino et al., 2013). The physical and
chemical characteristics of the waste, such as moisture, volatile solid, nutrient con-
tent, and particle size affect the biogas production and process stability. Cho et al.
(1995) determined the methane yields of different FW over 28 days at 37°C, and
found 482, 294, 277, and 472 mL/g VS for cooked meat, boiled rice, fresh cabbage,
and mixed FW, with 82%, 72%, 73%, and 86% efficiency, respectively, based on
elemental compositions of raw materials.
1.4.1 Single-Stage Anaerobic Digestion

The process configuration is very important for the efficiency of methane produc-
tion process. Single-stage anaerobic digestion process has been widely employed for
municipal solid waste treatment. As all of the reactions (hydrolysis, acidogenesis,
acetogenesis, and methanogenesis) take place simultaneously in a single reactor, the
system encounters less frequent technical failures and has a smaller investment cost
(Forster-Carneiro et al., 2008). The anaerobic digestion can be wet or dry; the former
uses the waste as received, while the latter needs to lower water content to about 12%
of total solid (Nasir et al., 2012). Compared to wet anaerobic digestion, dry anaerobic
digestion provides lower methane production and VS reduction due to the volatile
fatty acid (VFA) transport limitation (Nagao et al., 2012). El-Mashad et al. (2008)
reported that a digester treating FW was not stable due to the VFA accumulation and
low pH, leading to low biogas production. On the other hand, the stability of single-
stage anaerobic digester for easily degradable FW is of concern (Lee et al., 1999).
1.4.2 Two-Stage Anaerobic Digestion

In contrast to single-stage anaerobic digestion, two-stage anaerobic digestion has
often been used for producing both hydrogen and methane in two separate reactors
(Chu et al., 2008). In such a system, fast-growing acidogens and hydrogen-producing
microorganisms are enriched for the production of hydrogen and volatile fatty acids
16
TABLE 1.5
Methane Production from Food Wastes
OLR Biogas CH4
OLR (kg Yield Yield
Process Duration HRT (kg VS/ COD/ (mL/g (mL/g Efficiency
Waste Microorganism Pretreatment Type Vessel Type (days) (days) m3d) m3d) VS) VS) %CH4 (VS, %) References
Fruit and Cow manure None Two stage Bioreactor 29 1 1–9 NR NR 530 70 95.1 Mtz.
vegetable with 0.5 L Viturtia
waste working vol. et al.
(1989)
FW Anaerobic SS Freeze drying Two stage UASB with 120 NR 1.04 7–9 NR 277–482 NR 90 Cho et al.
of waste 8 L working (1995)
vol.
FW Anaerobic SS None Two stage Continuous 90 NR 7.9 NR NR 440 70 70 Lee et al.
pilot scale (1999)
5 tons/d
capacity
Fruit and Anaerobic SS None Single Serum bottles 100 Batch NA NA NR 180–732 NR NR Gunaseelan
vegetable stage with 135 mL (2004)
waste vol.
FW and Anaerobic SS None Single Semi 250 13 2.43 4.71 NR 321 64.4 55.8 Heo et al.
activated stage continuous (2004)
sludge reactor with
3.5 L
working vol.
(Continued)
OLR Biogas CH4
OLR (kg Yield Yield
Potato Anaerobic SS None Two stage Packed bed 38 NR NR 1–3 NR 390 82 NR Parawira
waste with 1 L et al.
working vol. (2005)
FW Anaerobic SS None Two stage Bioreactor 60 20 8 NR NR NR 68.8 86.4 Youn and
with 12 L Shin
working vol. (2005)
FW Bacteria None Single 3 stage semi 30 12 NR NR NR NR 67.4 NR Kim et al.
isolated from stage continuous (2006b)

landfill soil with 8 L
and cow working vol.
manure
FW Anaerobic SS None Single Batch 28 10–28 NA NA 600 440 73 81 Zhang et al.
stage (2007)
FW SS None Two stage CSTR with 150 5 6.6 16.3 NR 464 80 88 Chu et al.
10 L working (2008)
vol.
FW Landfill soil None Single Batch 5 L 60 20–60 NR NR 490 220 NR NR Forster-
and cow stage Carneiro
manure et al.
(2008)
(Continued)
17
18

OLR Biogas CH4
OLR (kg Yield Yield
FW Bacteria and None Three UASB with NR 12 54.5 ND ND 254 68 90.1 Kim et al.
sludge from stage 4800L (2008b)
various working vol.
sources
FW SS None Two stage Bioreactor 200 1–27 NR 15 578 520 90 NR Park et al.
with 4.5 L (2008)
working vol.
FW SS LAB Two stage Bioreactor 98 7 NR NR 850 434 51 NR Koike et al.
pretreatment with 5 L (2009)
and SsF working vol.
FW No addition None Two stage Rotating drum 30 SRT 4.61 NR 769 546 71.5 82.2 Wang and
with 200 L 26.7 h Zhao
working vol. (2009)
FW SS Heat Two stage UASB with 60 3.9–6.4 NR NR NR NR 80 80 Lee et al.
pretreatment 2.3 L (2010a)
(100°C working vol.
30 min)
FW SS None Two stage Gas sparging 96 15.4 NR 4.16 NR NR 65 88.1 Lee and
type reactor Chung
with 40 L (2010)
working vol.
(Continued)
Another random document with
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form communities consisting at times of a countless number of
individuals; but it has not been thoroughly ascertained whether these
are the produce of a single queen, as in the case of the hive-bee, or
whether there may be more than one egg-producer in each
community. The late F. Smith thought the former of these alternatives
would prove to be correct. These mosquito-bees are frequently
spoken of as stingless bees, but this is not quite correct, for although
they do not sting, von Ihering[34] says that all the essential elements
of the sting are present, the pointed or penetrating part of the
apparatus being stunted.
It would serve no useful purpose to attempt to construct the social

history of these stingless bees from the numerous brief scattered
accounts in entomological literature, for they refer to different
species; it is, however, positively stated by Smith on the authority of
Peckolt[35] that Trigona mosquito sends off swarms after the manner
of the hive-bee in this country, and that after searching six hives only
one royal female could be found in each.
Fig. 24.—Melipona sp. ♀. Amazons.
The nests of many of these little bees are rich in honey, and they
have a host of enemies from man and monkeys downwards; and as
they do not defend themselves by stinging, it might be supposed
they would have but a poor time of it. From the accounts that have
been published we may, however, gather that they are rich in
devices for the protection of their nests, and for the exclusion of
intruders. Bates has given some particulars as to Melipona interrupta
(fasciculata); it is about one-third shorter than the hive-bee, and its
colonies are composed of an immense number of individuals. The
workers are usually occupied in gathering pollen; but they also
collect clay in a similar manner, and convey it to the nest, where it is
used for building a wall to complete the fortification of the nest, which
is placed either in a suitable bank, or in a trunk of a tree; in either
situation it is completely built in with clay. A nest which Bates saw
opened contained about two quarts of pleasantly-tasted liquid honey.
Forty-five species of these little bees were found in different parts of
the Amazons Valley, the largest kind being half an inch in length, the
smallest very minute, not more than one-twelfth of an inch. These
little creatures are thus masons as well as workers in wax and resin,
and they are also gatherers of nectar, pollen, and resin.
According to Gosse, one of these bees is well known in Jamaica,

where they are called "Angelitos," in consequence of their not
stinging people. He observed a nest of this bee in a tree, and found it
to be much infested by black ants anxious to obtain entrance to it;
three bees, however, stood sentinel in the entrance, so as to
completely block it and keep out intruders, but the middle bee moved
on one side out of the way directly one of its fellows wished to come
in or out of the nest. The honey accumulated by this species is kept
in clusters of cups about the size of a pigeon's egg, at the bottom of
the hive and away from the brood-cells. The queen or mother-bee is
lighter in colour than the others, and has the hind body twice the
length of theirs.
Hockings[36] has given us some details as to the natural history of

two of these bees that inhabit Australia, where they are called
"Karbi" and "Kootchar," the first being, it is supposed, Trigona
carbonaria, Smith: it is usually about three-sixteenths of an inch in
length, the queen, when fully developed, being nearly twice that
length. The comb is built in a most peculiar form, being, it is said, in
the shape of a spiral staircase, and tapering towards the ends:
honey-pots and pollen are constructed for the storage of food. The
comb is encased in wax, and outside it a labyrinth of waxen
passages is formed. The entrance to the colony is guarded by a line
of bees who inspect every one that arrives, and it is surprising to see
how soon a stranger is discovered and pounced upon before it has
time even to alight; the intruder, when caught, is held by several
bees, who put it on the rack by holding and stretching out its limbs to
their full extent, retaining it in this position for as long as an hour, by
which time the unfortunate prisoner is usually dead. These bees, as
well as many other allied species, fight desperately with their
mandibles, and are apparently of a very fierce disposition. The other
species, called "Kootchar," is said to produce a very large number of
drones, and the habits and dispositions of the bees differ
considerably from those of the "Karbi": the entrance to their hive is
guarded by a pipe of propolis (a sort of resinous wax) about an inch
in length, having an exceedingly sticky outer edge, and it is by this
pipe alone that access to the interior can be gained. At night the
entrance is closed by numerous minute globules of semi-fluid gum
placed against it, thus forming a thin wall full of air-holes. The
colonies of "Kootchar" can be united by taking away a queen and
then packing her brood-nest, bees and all, against that of the colony
it is to be joined to. This cannot be done with the "Karbi." The
account given by Mr. Hockings contains a great many other
interesting details, and there can be no doubt that a full account of
the natural history of these Insects would be very instructive.
Fritz Müller has recorded a singular case bearing on the instinct of

these social Insects: he says that a nest of a small Trigona was built
in a hollow tree, and that as a consequence of the irregularity of the
hole the bees were obliged to give a very irregular shape to their
combs of honey. These bees were captured and put in a spacious
box (presumably together with the irregular comb, but this he
unfortunately does not mention): after a year, "when perhaps not a
single bee survived of those which had come from the canella tree,"
they still continued to build irregular combs, though quite regular
combs were built by several other communities of the same species
that he had kept. These bees, he also tells us, do not use pure wax
for the construction of their combs, but mix it with resin or gum that
gives it a peculiar odour and appearance. He captured two
communities of a common Melipona, one of which had the combs
made of dark reddish brown, the other of pale yellowish brown, wax,
and in captivity in a distant locality each of the two communities
continued to form its comb in the same way, thus showing the
continuity that prevails in these cases as long as circumstances
permit. Müller thinks this due to imitation, but it seems at least as
probable that it is due to perception of the properties of the nest. The
nest has a certain colour that the worker-bee matches.
Several species of the Melipona and Trigona were imported from

Brazil to France, and kept there for some time in captivity by M.
Drory. Girard has published[37] some details as to these colonies,
and is of opinion that some of them indicate an intelligence or instinct
superior to that of the honey-bee. The queen-bee of M. scutellaris
seems to display more intelligence than the corresponding sex of A.
mellifica. The mode of feeding the larvae apparently differs from that
of A. mellifica, a provision of pollen being first placed in the cell, then
some honey; when sufficient food for the whole consumption of a
larva is accumulated the queen deposits an egg in the cell, which is
at once completely closed by the worker. The interior of the abode of
these bees is quite dark, only a very small orifice being left, and in
this a sentinel is constantly on the alert. The same writer states that
Trigona crassipes has the very peculiar habit of always locating its
brood-comb in the nest of a species of Termes.
The honey-bee, Apis mellifica (Fig. 6), is considered the highest form
attained by the Anthophilous division of the Hymenoptera. The
differentiation of the three forms, male, female, and worker, is here
carried to a greater degree of perfection than in the other bees. The
drones are the males; the individuals we see gathering honey are
always workers, neither the male nor the female in this species
taking any part in procuring food for themselves or for the colony. In
addition to this the colonies formed may be described as permanent:
they do not come to an end at the close of one season, and
provision is made for the formation of a new colony while the old one
still persists, by means of a peculiar process called swarming. The
life-history of Apis mellifica and its anatomy and physiology have
been discussed in a whole library of works, and we need only notice
the chief features. When a swarm of bees leaves a hive it consists of
the queen-bee or female, and a number of workers, these latter
being, in fact, the surplus population that has been produced in the
hive. The swarm is not a nuptial flight, as is often supposed, but an
act of emigration. When this swarm has been housed, the bees
commence operations in their new quarters, by secreting wax; they
are enabled to do this by having consumed much saccharine food;
the wax is produced by means of glands in the hind-body over the
inner faces of the ventral plates of the abdominal rings, and it makes
its appearance there, after passing from the interior of the body
through some peculiar membranes on the ventral segments, in the
form of thin projecting plates. These the bee takes off with an
apparatus on the hind pair of legs and applies, after working up with
the mandibles, to form the cells in which young ones are to be
reared and food stored. A large number of bees working in common
thus produce the regular and beautiful structure known as the comb;
the queen afterwards lays an egg in each cell, and as these soon
hatch, great labour is thrown on the workers, which have then to
feed the young; this they do by eating honey and pollen, which,
being formed into a sort of pap by a portion of their digestive organs,
is then regurgitated and given to the young, a quantity of it being
placed in the cell, so that the larva is bathed by it, and possibly may
absorb the food by the skin as well as the mouth. When the colony is
in good progress and young bees emerge, these act as nurses, the
older ones cease to prepare food and act as foragers, bringing in
honey and pollen which are each stored in separate cells. The larva
in the cell increases its size and sheds a very delicate skin several
times; when the larva has reached its full size no more food is
supplied, but the worker-bees seal up the cell by means of a cover
formed of pollen and wax, in such a manner as to be pervious to air:
sealed up in the cell the larva spins a cocoon for itself, remains
therein for a little time as a larva, then changes to a pupa, and
thereafter bites its way out through the cover of the cell, and appears
for the first time as a new being in the form of a worker-bee; the
whole process of development from the egg-state to the perfect
condition of the worker-bee occupies about three weeks.
When the denizens of a hive are about to produce another queen,
one or more royal cells are formed; these are much larger than the
ordinary worker-cells, and of a quite different form. In this cell is
placed an egg, not differing in any respect from the egg that, if
placed in an ordinary cell, produces a worker; when the egg has
produced a larva this is tended with great care and fed throughout its
life with royal jelly. This food appears to be the same as that supplied
to an ordinary worker-larva when it is first hatched; but there is this
difference, that whereas the worker-larva is weaned, and supplied,
after the first period of its existence, with food consisting largely of
honey, pollen and water, the queen-larva is supplied with the pap or
royal jelly until it is full grown. Some difference of opinion exists as to
this royal jelly, some thinking that it is a different substance from
what the workers are fed with; and it is by no means improbable that
there may be some difference in the secretion of the glands that
furnish a part of the material composing the pap. The queen is
produced more rapidly than workers are, about sixteen days being
occupied in the process of her development. Only one queen is
allowed in a hive at a time; so that when several queen-cells are
formed, and queen-larvae nurtured in them, the first one that is
developed into a perfect queen goes round and stings the royal
nymphs to death while they are still in their cells. The production of
drones is supposed to depend chiefly on the nature of the egg laid
by the queen; it being considered that an unfertilised egg is
deposited for this purpose. There is still some doubt on this point,
however. Though there is no doubt that drones are produced in great
numbers from unfertilised eggs, yet there is not evidence that they
cannot also be produced from fertilised eggs.[38] The drone-cells are
somewhat larger than the ordinary worker-cells, but this is probably
not of much import, and it is said that the larvae intended to produce
drones receive a greater proportion of pap than worker-larvae do:
about twenty-four days are required to produce a drone from the
egg.
From this sketch it will be seen that the production of the worker (or
third sex, as it is improperly called, the workers being really females
atrophied in some points and specially developed in others) is
dependent on the social life, in so far at any rate as the special
feeding is concerned. There is good reason for supposing that A.
mellifica has been kept in a state of domestication or captivity for an
enormous period of time; and this condition has probably led to an
increase of its natural peculiarities, or perhaps we should say to a
change in them to suit a life of confinement. This is certainly the case
in regard to swarming, for this process takes place with comparative
irregularity in Apis mellifica in a wild condition. The killing of
superfluous queens is also probably a phenomenon of captivity, for it
varies even now in accordance with the numbers of the colony. It is
interesting to notice that in confinement when a swarm goes from the
hive it is the old queen that accompanies it, and this swarm as a rule
settles down near the old hive, so that the queen-bee being already
fertilised, the new swarm and its subsequent increase are nothing
but a division of the old hive, the total products of the two having but
a single father and mother. When a second swarm goes off from a
hive it is accompanied by a young queen, who frequently, perhaps,
in the majority of cases, is unfertilised; this swarm is apt to fly for
long distances, so that the probability of cross-fertilisation is greatly
increased, as the fertilisation of the young new queen is effected
during a solitary flight she makes after the colony has settled down.
But in a state of nature the colonies do not send off swarms every
year or once a year, but increase to an enormous extent, going for
years without swarming, and then when their home is really filled up
send off, it may be presumed, a number of swarms in one year. Thus
the phenomena of bee-life in a wild condition differ considerably from
those we see in artificial confinement. And this difference is probably
greatly accentuated by the action of parasites, the proportions of
which to their guests are in a state of nature liable to become very
great; as we have seen to be the case in Bombus.
Under these circumstances it is not a matter for surprise when we

find that the honey-bee has formed distinct races analogous to those
that exist in the case of the domesticated vertebrate animals. The
knowledge of these races is, however, at present very little
advanced, and is complicated by the fact that only imperfect
information exists as to the true species of the genus Apis. There is
a bee very like our common honey-bee found in southern Europe
called A. ligustica; this is certainly a variety of A. mellifica, and the
same remark applies to a bee found in Egypt, and called A. fasciata.
This gives the honey-bee a very wide distribution, extending possibly
over the whole of the palaearctic region: besides this, the species
has been introduced into various other parts of the world.
According to Karsch the honey-bee shows in Germany several

varieties, all of which belong to the northern form, which may be
spoken of as the A. domestica of Ray; the A. ligustica and A. fasciata
form as we have said distinct races, and it is a remarkable fact that
these races remain distinct even when imported into other climates;
though for how long a period of time this remains true there is very
little evidence to show. The northern form, A. domestica, is now
found in very widely separated parts of the world, in some of which it
is wild; Smith mentions it as occurring in the West India islands,
throughout the North American continent as far south as Mexico,
even in Central and Southern Africa, and in Australia and New
Zealand. The var. ligustica has been found also at the Cape of Good
Hope. The other species known of the genus Apis all belong to the
Old World, so that there is very little doubt that A. mellifica is also a
true native of the eastern hemisphere, and its original home may
possibly have been not far from the shores of the eastern portion of
the Mediterranean sea. Seven or eight other species of Apis are
known, all but one of which occur in Asia, extending as far as Timor
and Celebes. The exceptional one, A. adansonii, occurs in tropical
Africa and in Madagascar. Gerstaecker thought these species might
be reduced to four, but Smith's statement that the males and even
the workers show good distinctive characters seems to be correct.
Very little is known as to the honey-bees of China and Japan.
The queen-bee greatly resembles the worker, but has the hind body
more elongated; she can, however, always be distinguished from the
worker by the absence of the beautiful transverse, comb-like series
of hairs on the inner side of the first joint of the hind foot, the planta,
as it is called by the bee-keeper: she has also no wax plates and
differs in important anatomical peculiarities. The male bee or drone
is very different, being of much broader, more robust build, and with
very large eyes that quite meet in the middle of the upper part of the
head: he also has the hind leg differently shaped. The form of this
limb enables the male of A. mellifica to be distinguished from the
corresponding sex of allied species of the genus.
Fig. 25.—Portions of hind-feet, 1, of male, 2, of worker, 3, of queen, of

the honey-bee; series on the left, outer faces; on the right, inner
faces. a, Tip of tibia: b, first joint; c, second joint of tarsus.
We are indebted to Horne for some particulars as to the habits of A.

dorsata, an allied East Indian species. He informs us that these bees
greatly disfigure buildings, such as the Taj Mahal at Agra, by
attaching their pendent combs to the marble arches, and are so
pertinacious that it is almost useless to destroy the nests. This bee is
said to be so savage in its disposition that it cannot be domesticated;
it attacks the sparingly clad Hindoos with great ferocity when they
disturb its nest. Notwithstanding its inclination and power to defend
its societies this Insect appears to be destroyed wholesale. Colonel
Ramsay failed to establish hives of it, because the Insects were
eaten up by lizards. The crested honey-buzzard carries off large
portions of the comb, and devours it on a branch of some tree near
by, quite regardless of the stings of the bees; while the fondness of
bears for the honey of the "Dingar," as this species is called, is well
known.
Note to P. 33: It has just been discovered that a most remarkable
symbiosis, with structural modification of the bee, exists between
the females of Xylocopa, of the Oriental sub-genus
Koptorthosoma, and certain Acarids. A special chamber, with a
small orifice for entry, exists in the abdomen of the bee, and in
this the Acari are lodged.—See Perkins, Ent. Mag. xxxv. 1899, p.
37.
Note to P. 80: referring to the habits of social wasps in warm

countries. The anticipation we ventured to indulge in is shown to
be correct by the recent observations of Von Ihering.[39] He
states that social wasps in Brazil may be divided into two great
groups by their habits, viz. 1. Summer communities, lasting for
one year, and founded annually by fertilised females that have
hibernated—example, Polistes; 2. Perennial communities,
founded by swarms after the fashion of bee colonies—examples,
Polybia, Chartergus.
Note to Vol. V. Pp. 545, 546: The development of Encyrtus

fuscicollis has now been studied by Marchal, who has
discovered the existence of embryonic dissociation. The chain of
embryos and the epithelial tube in which they are placed, are
formed as follows: the Encyrtus deposits an egg in the interior of
the egg of the Hyponomeuta. This does not kill the egg of the
Lepidopteron, but becomes included in the resulting caterpillar.
The amnion of the Chalcid egg lengthens, and forms the
epithelial tube; while the cells within it become dissociated in
such a way as to give rise to a chain of embryos, instead of a
single embryo.—C.R. Ac. Paris, cxxvi. 1898, p. 662, and
translation in Ann. Nat. Hist. (7), ii. 1898, p. 28.
CHAPTER II
HYMENOPTERA ACULEATA CONTINUED—DIVISION II. DIPLOPTERA OR

WASPS—EUMENIDAE, SOLITARY TRUE WASPS—VESPIDAE, SOCIAL
WASPS—MASARIDAE
Division II. Diploptera—Wasps.
Anterior wings longitudinally plicate in repose; the pronotum

extending back, so as to form on each side an angle reposing on
the tegula; the basal segments of the hind body not bearing
nodes or scales; the hind tarsi formed for simple walking. The
species either solitary or social in their habits; some existing in
three forms, males, females, and workers.
Fig. 26—Upper aspect of pronotum and mesonotum of a wasp,

Eumenes coarctata. a, Angle of pronotum; b, tegula; c, base of
wing; d, mesonotum.
This division of Hymenoptera includes the true wasps, but not the
fossorial wasps. The name applied to it has been suggested by the
fact that the front wings become doubled in the long direction when
at rest, so as to make them appear narrower than in most other
Aculeata (Fig. 27). This character is unimportant in function so far as
we know,[40] and it is not quite constant in the division, since some of
the Masaridae do not exhibit it. The character reappears outside the
Diploptera in the genus Leucospis—a member of the Chalcididae in
the parasitic series of Hymenoptera—the species of which greatly
resemble wasps in coloration. A better character is that furnished by
the well-marked angle, formed by the pronotum on the dorsal part
(Fig. 26). By a glance at this part a Diplopterous Insect can always
be readily distinguished.
Three families are at present distinguished in the Diploptera, viz.

Eumenidae, Vespidae and Masaridae. We anticipate that Eumenidae
and Vespidae will ultimately be found to constitute but one family.
Fam. 1. Eumenidae—Solitary True Wasps.
Claws of the feet toothed or bifid; middle tibiae with only one
spur at tip. Social assemblages are not formed, and there is no
worker-caste, the duties of nest-construction, etc., being
performed solely by the female.
The Eumenidae, or solitary wasps, are very little noticed by the

ordinary observer, but they are nevertheless more numerous than
the social Vespidae, about 800 species being known. In Britain we
have sixteen species of the solitary, as against seven of the social
wasps. The Eumenidae exhibit a considerable diversity in form and
structure; some of them have the pedicel at the base of the abdomen
very elongate, while in others this is so short as to be imperceptible
in the ordinary position of the body. A repetition of similar differences
of form occurs in the social wasps, so that notwithstanding the
difference in habits there seems to be no satisfactory way of
distinguishing the members of the two families except by the
structure of the claws and tibial spurs.
Fig. 27.—Eumenes flavopicta ♀. Burma. The wings on the left in the

position of repose, to show folding.
Fabre has sketched the habits of a species of Eumenes, probably E.
pomiformis. This Eumenes constructs with clay a small vase-like
earthenware vessel, in the walls of which small stones are
embedded (like Fig. 28, B). This it fills with food for the young. The
food consists of caterpillars to the number of fourteen or sixteen for
each nest. These caterpillars are believed to be stung by the parent-
wasp (as is the case in the fossorial Hymenoptera), but complete
evidence of this does not seem to be extant, and if it be so, the
stinging does not completely deprive the caterpillars of the capacity
of movement, for they possess the power of using their mandibles
and of making strokes, or kicking with the posterior part of the body.
It is clear that if the delicate egg of the Eumenes or the delicate larva
that issues from it were placed in the midst of a mass of this kind, it
would probably suffer destruction; therefore, to prevent this, the egg
is not placed among the caterpillars, but is suspended from the
dome covering the nest by a delicate thread rivalling in fineness the
web of the spider, and being above the mass of food it is safe. When
the young larva leaves the egg it still makes use of the shell as its
habitation, and eats its first meals from the vantage-point of this
suspension; although the mass of the food grows less by
consumption, the little larva is still enabled to reach it by the fact that
the egg-shell splits up to a sort of ribbon, and thus adds to the length
of the suspensory thread, of which it is the terminal portion. Finally
the heap of caterpillars shrinks so much that it cannot be reached by
the larva even with the aid of the augmented length of the
suspensory thread; by this time, however, the little creature has so
much increased in size and strength that it is able to take its place
amongst the food without danger of being crushed by the mass, and
it afterwards completes its metamorphosis in the usual manner.
Fig. 28—Nidification of solitary wasps: section through nest, A, of
Odynerus reniformis; B, of Eumenes arbustorum. a, The
suspended egg of the wasp; b, the stored caterpillars. (After
André.)
It is known that other species of Eumenes construct vase-like nests;

E. unguiculata, however, according to an imperfect account given by
Perris, makes with earth a closed nest of irregular shape, containing
three cells in one mass. The saliva of these builders has the power
of acting as a cement, and of forming with the clay a very
impenetrable material. One species, E. coarctata, L. of this genus
occurs in Britain. The clay nests (Fig. 29) of this Insect are often
attached to the twigs of shrubs, while those of the two species
previously mentioned are usually placed on objects that offer a large
surface for fixing the foundations to, such as walls. According to
Goureau the larva of this species forms in one corner of its little
abode, separated by a partition, a sort of dust-heap in which it
accumulates the various débris resulting from the consumption of its
stores.
Eumenes conica, according to Horne, constructs in Hindostan clay-

nests with very delicate walls. This species provisions its nest with
ten or twelve green caterpillars; on one occasion this observer took
from one cell eight green caterpillars and one black. It is much
attacked by parasites owing, it is thought, to the delicacy of the walls
of the cells, which are easily pierced; from one group of five cells two
specimens only of the Eumenes were reared.
Fig. 29—Nest of Eumenes coarctata: A, the nest attached to wood; B,

detached, showing the larva. a, the larva; b, the partition of the
cell. (After André.)
Odynerus, with numerous sub-genera, the names of which are often
used as those of distinct genera, includes the larger part of the
solitary wasps; it is very widely distributed over the earth, and is
represented by many peculiar species even in the isolated
Archipelago of Hawaii; in Britain we have about fifteen species of the
genus. The Odynerus are less accomplished architects than the
species of Eumenes, and usually play the more humble parts of
adapters and repairers; they live either in holes in walls, or in posts
or other woodwork, or in burrows in the earth, or in stems of plants.
Several species of the sub-genus Hoplopus have the remarkable
habit of constructing burrows in sandy ground, and forming at their
entry a curvate, freely projecting tube placed at right angles to the
main burrow, and formed of the grains of sand brought out by the
Insect during excavation and cemented together. The habits of one
such species were described by Réaumur, of another by Dufour; and
recently Fabre has added to the accounts of these naturalists some
important information drawn from his own observations on O.
reniformis.
Fig. 30.—Odynerus antilope ♀. Britain.
This Insect provisions its cell with small caterpillars to the number of
twenty or upwards (Fig. 28, A.) The egg is deposited before the nest
is stocked with food; it is suspended in such a manner that the
suspensory thread allows the egg to reach well down towards the
bottom of the cell. The caterpillars placed as food in the nest are all
curled up, each forming a ring approximately adapted to the calibre
of the cell. Fabre believes these caterpillars to be partly stupefied by
stinging, but the act has not been observed either by himself,
Réaumur, or Dufour. The first caterpillar is eaten by the wasp-larva
from its point of suspension; after this first meal has been made the
larva is supposed to undergo a change of skin; it then abandons the
assistance of the suspensory thread, taking up a position in the
vacant chamber at the end of the cell and drawing the caterpillars to
itself one by one. This arrangement permits the caterpillars to be
consumed in the order in which they were placed in the cell, so that
the one that is weakest on account of its longer period of starvation
is first devoured. Fabre thinks all the above points are essential to
the successful development of this wasp-larva, the suspension
protecting the egg and the young larva from destruction by pressure
or movement of the caterpillars, while the position of the larva when
it leaves the thread and takes its place on the floor of the cell
ensures its consuming the food in the order of introduction; besides
this the caterpillars used are of a proper size and of a species the
individuals of which have the habit of rolling themselves up in a ring;
while, as the calibre of the tube is but small, they are unable to
straighten themselves and move about, so that their consumption in
proper order is assured. Some interesting points in the habits of an
allied species, O. (Pterocheilus) spinipes have been observed by
Verhoeff; the facts as regards the construction and provisioning of
the cell are almost the same as in O. reniformis. The species of
Odynerus are very subject to the attacks of parasites, and are, it is
well known, destroyed to an enormous extent by Chrysididae.
Verhoeff says that the wasp in question supplied food much infested
by entoparasites; further, that a fly, Argyromoeba sinuata, takes
advantage of the habit of the Odynerus of leaving its nest open
during the process of provisioning, and deposits also an egg in the
nest; the Odynerus seems, however, to have no power of
discovering the fact, or more probably has no knowledge of its
meaning, and so concludes the work of closing the cell in the usual
way; the egg of the Argyromoeba hatches, and the maggot produced
feeds on the caterpillars the wasp intended for its own offspring.
Verhoeff observed that the egg of the wasp-larva is destroyed, but
he does not know whether this was done by the mother
Argyromoeba or by the larva hatched from her egg. Fabre's
observations on allied species of Diptera render it, however, highly
probable that the destruction is effected by the young fly-larva and
not by the mother-fly.
Mr. R. C. L. Perkins once observed several individuals of our British
O. callosus forming their nests in a clay bank, and provisioning them
with larvae, nearly all of which were parasitised, and that to such an
extent as to be evident both to the eye and the touch. In a few days
after the wasps' eggs were laid, swarms of the minute parasites
emerged and left no food for the Odynerus. Curiously, as it would
seem, certain of the parasitised and stored-up larvae attempted (as
parasitised larvae not infrequently do), to pupate. From which, as Mr.
Perkins remarks, we may infer that (owing to distortion) the act of
paralysing by the wasp had been ineffectual. Mr. Perkins has also
observed that some of the numerous species of Hawaiian Odynerus
make a single mud-cell, very like the pot of an Eumenes, but
cylindrical instead of spherical. This little vessel is often placed in a
leaf that a spider curls up; young molluscs of the genus Achatinella
also avail themselves of this shelter, so that a curious colony is
formed, consisting of the Odynerus in its pot, of masses of the young
spiders, and of the little molluscs.
Horne has recorded that the East Indian O. punctum is fond of

availing itself of holes in door-posts where large screws have been;
after the hole has been filled with provisions, the orifice is covered
over level with the surface of the wood so that it eludes human
observation. It is nevertheless discovered by an Ichneumon-fly which
pierces the covering with its ovipositor and deposits an egg within.
The genus Abispa is peculiar to Australia and includes some very

fine solitary wasps, having somewhat the appearance of very large
Odynerus: these Insects construct a beautiful nest with a projecting
funnel-shaped entrance, and of so large a size that it might pass for
the habitation of a colony of social wasps; it appears, however, that
this large nest is really formed by a single female.
The species of the genus Rhygchium are also of insecticide habits,

and appear to prefer the stems of pithy plants as the nidus for the
development of the generation that is to follow them. Lichtenstein
says that a female of the European R. oculatum forms fifteen to
twenty cells in such a situation, and destroys 150 to 200 caterpillars,
and he suggests that, as it is easy to encourage these wasps to nest
in a suitable spot, we should utilise them to free our gardens from
caterpillars, as we do cats to clear the mice from our apartments.
The East Indian R. carnaticum seems to have very similar habits to

its European congener, adapting for its use the hollow stems of
bamboos. Horne has recorded a case in which a female of this
species took possession of a stem in which a bee, Megachile lanata,
had already constructed two cells; it first formed a partition of mud
over the spot occupied by the bee, this partition being similar to that
which it makes use of for separating the spaces intended for its own
young. This species stores caterpillars for the benefit of its larvae,
and this is also the case with another Eastern species, R. nitidulum.
This latter Insect, however, does not nidificate in the stems of plants,
but constructs clay cells similar to those of Eumenes, and fixes them
firmly to wood. Rhygchium brunneum is said by Sir Richard Owen to
obliterate hieroglyphic inscriptions in Egypt by its habit of building
mud nests amongst them. An individual of this wasp was found by
Dr. Birch when unrolling a mummy—"There being every reason to
believe that the Insect had remained in the position in which it was
found ever since the last rites were paid to the ancient Egyptian."
Fam. 2. Vespidae—Social Wasps.
Claws of the feet simple, neither toothed nor bifid, middle tibiae
with two spurs at the tip. Insects living in societies, forming a
common dwelling of a papery or card-like material; each
generation consists of males and females and of workers—
imperfect females—that assist the reproductive female by
carrying on the industrial occupations.
The anterior wing possesses four submarginal cells, as in the

Eumenidae. The attention of entomologists has been more directed
to the habits and architecture than to the taxonomy of these Insects,
so that the external structure of the Insects themselves has not been
so minutely or extensively scrutinised as is desirable; de Saussure,
the most important authority, bases his classification of the Insects
themselves on the nature of the nests they form. These habitations
consist of an envelope, protecting cells similar in form to the comb of
the honey-bee, but there is this important difference between the
two, that while the bee forms its comb of wax that it secretes, the
wasps make use of paper or card that they form from fragments of
vegetable tissue,—more particularly woody fibre—amalgamated by
means of cement secreted by glands; the vegetable fragments are
obtained by means of the mandibles, the front legs playing a much
less important part in the economy of the Vespidæ than they do in
that of the bees and fossorial Hymenoptera.
In most of the nests of Vespidæ the comb is placed in stages or

stories one above the other, and separated by an intervening space,
but in many cases there is only one mass of comb. It is the rule that,
when the cells of the comb are only partially formed, eggs are
deposited in them, and that the larva resulting from the egg is fed
and tended by the mother, or by her assistants, the workers; as the
larvae grow, the cells are increased in correspondence with the size
of the larva; the subsequent metamorphosis to pupa and imago
taking place in the cells after they have been entirely closed. The
food supplied is of a varied nature according to the species, being
either animal or vegetable, or both.
Fig. 31—Section of the subterranean nest of the common wasp, Vespa

germanica, in position. (After Janet.) a, One of the chambers of an
ant's nest, Lasius flavus, placed above the wasps' nest; b, root to
which the first attachment of the nest was made; c, secondary
attachments; d, the first-made attachment; e, a flint within the
envelopes of the nest; f, the chief suspensory pillar of the second
layer of comb; g, lateral galleries; h, one of the secondary pillars
of suspension between two layers of comb; i, the layers of wasp-
paper forming the envelope of nest; j, vacant space round the
nest; k, flints that fell to the bottom during the work of excavation;
l, numerous larvae of a fly, Pegomyia inanis (?) placed vertically in
ground beneath the nest; m1 to m7, the layers of comb, in m2 the
cells are indicated, in m8 (above the main figure) the arrangement
of the three cells forming the commencement of the new layer of
comb, m7, is shown; n, gallery of access from surface; o, burrow
of a mole; p, interval of 90 mm. between top of nest and surface;
q, height of the nest, 163 mm.
Although the nests of the social wasps are very elaborate

constructions, yet they serve the purposes of the Insects for only a
single season. This is certainly the case in our own country. Here
each nest is commenced by a single female or queen; she at first
performs unaided all the duties for the inauguration of the colony;
she lays the foundation of the cells, deposits the eggs in them, feeds
the young, and thus rears a brood of workers that at once assist her,
and for the future relieve her of a considerable portion of her former
occupations; the nest is by them added to and increased, till the cold
weather of the autumn is at hand; at this time many males and
females are produced; the cold weather either destroys the
inhabitants of the nest, or reduces their vitality so that it is impossible
for them to pursue successfully the avocations necessary for their
subsistence, and they succumb to adversity. The young females,
however, hibernate, and each one that lives through the winter is the
potential founder of a new nest in the way we have already
described. It might be supposed that in tropical countries where no
cold season occurs the phenomena would be different, that the
colonies would be permanent, and that the nests would be inhabited
until they were worn out. De Saussure, however, informs us that this
is not the case, but that in the tropics also the colonies die off
annually. "The nests are abandoned," he says, "without it being

Biofuels From Food Waste: Applications of Saccharification Using Fungal Solid State Fermentation 1st Edition Antoine Prandota Trzcinski

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Biofuels From Food Waste: Applications of Saccharification Using Fungal Solid State Fermentation 1st Edition Antoine Prandota Trzcinski

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Biofuels from Food Waste: Applications

of Saccharification using Fungal Solid

Solid State Fermentation Research and Industrial

Advances in Solid Biofuels Elias Christoforou

Using Energy Crops for Biofuels or Food The Choice

Solid State Properties From Bulk to Nano 1st Edition

Handbook of Food Bioengineering Volume 3 Soft Chemistry

Solid State Chemistry and its Applications Student

Introductory Solid State Physics with MATLAB

Solid State Chemistry 1st Edition Frank Hoffmann

© 2018 by Taylor & Francis Group, LLC

No claim to original U.S. Government works

Printed on acid-free paper

International Standard Book Number-13: 978-1-138-09372-0 (Hardback)

Library of Congress Cataloging-in-Publication Data

Names: Trzcinski, Antoine Prandota, author.

Visit the Taylor & Francis Web site at

Chapter 1 Bioconversion of Food Wastes to Energy.............................................1

Chapter 2 Platform Chemical Production from Food Wastes.............................25

Chapter 3 Enzyme Production from Food Wastes............................................... 49

Chapter 4 Enhanced Glucoamylase Production by Aspergillus awamori

Chapter 5 Enhancing the Hydrolysis and Methane Production Potential of

5.2.4 Anaerobic Digestion of Enzymatically

b Melikoglu et al. (2013b)

e Noor et al. (2013)

1.2 ETHANOL PRODUCTION

maltose syrups, and fructose) can be produced in saccharification process, whereas

1.2.3 Process Configurations

1.2.4 Other Strategies to Improve Ethanol Yield

working vol. KA4 (2008c)

TABLE 1.3 (Continued)

1.2.5 Large-Scale Ethanol Production from Food Waste

1.3 HYDROGEN PRODUCTION

TABLE 1.4 (Continued)

1.3.3 Process Configurations

1.4 METHANE PRODUCTION

1.4.1 Single-Stage Anaerobic Digestion

1.4.2 Two-Stage Anaerobic Digestion

isolated from stage continuous (2006b)

TABLE 1.5 (Continued)

It would serve no useful purpose to attempt to construct the social

Fig. 24.—Melipona sp. ♀. Amazons.

According to Gosse, one of these bees is well known in Jamaica,

Hockings[36] has given us some details as to the natural history of

Fritz Müller has recorded a singular case bearing on the instinct of

Several species of the Melipona and Trigona were imported from

Under these circumstances it is not a matter for surprise when we

According to Karsch the honey-bee shows in Germany several

Fig. 25.—Portions of hind-feet, 1, of male, 2, of worker, 3, of queen, of

We are indebted to Horne for some particulars as to the habits of A.

Note to P. 80: referring to the habits of social wasps in warm

Note to Vol. V. Pp. 545, 546: The development of Encyrtus

HYMENOPTERA ACULEATA CONTINUED—DIVISION II. DIPLOPTERA OR

Division II. Diploptera—Wasps.

Anterior wings longitudinally plicate in repose; the pronotum

Fig. 26—Upper aspect of pronotum and mesonotum of a wasp,

Three families are at present distinguished in the Diploptera, viz.

Fam. 1. Eumenidae—Solitary True Wasps.

The Eumenidae, or solitary wasps, are very little noticed by the

Fig. 27.—Eumenes flavopicta ♀. Burma. The wings on the left in the

It is known that other species of Eumenes construct vase-like nests;

Eumenes conica, according to Horne, constructs in Hindostan clay-

Fig. 29—Nest of Eumenes coarctata: A, the nest attached to wood; B,