Renewable and Sustainable Energy Reviews 139 (2021) 110691

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Renewable and Sustainable Energy Reviews

journal homepage: http://www.elsevier.com/locate/rser

Biomass for a sustainable bioeconomy: An overview of world biomass

production and utilization
Mohammed Antar a, 1, Dongmei Lyu a, 1, Mahtab Nazari a, 1, Ateeq Shah a, 1, Xiaomin Zhou a, b, 1,
Donald L. Smith a, 1, *
a
Department of Plant Science, McGill University, Sainte-Anne-de-Bellevue, Quebec, Canada
b
BiofuelNet Canada Network, Sainte-Anne-de-Bellevue, Quebec, Canada

A R T I C L E I N F O A B S T R A C T

Keywords: Given both the ongoing expansion of world population and development of climate change conditions, it is
Biomass production increasingly imperative to develop and deploy sustainable biomass production methods to allow establishment of
Bioeconomy a flourishing and sustainable bioeconomy. Green technologies, including biofuels and bioproducts, are among
Biofuel
the most effective strategies for decreasing greenhouse gas emissions and global warming, while meeting
Renewable energy
humanity’s energy requirements. Biomass now provides a measure of energy to many countries, however sup­
Beneficial microorganism
Genome editing porting technologies are not widely accepted, largely because of low returns for biomass producers. This paper
Agronomic practices provides an overview of world biomass production and utilization. It also indicates potential approaches for
Microbe-to-plant signal compounds enhancing biomass production: agronomic practices, associated microorganisms, genome editing, selection of
optimal technologies, best combination approaches for feeding global human and animal populations, while,
decreasing greenhouse gas emissions and replacing demand for fossil energy with bioenergy. A more novel
concept is proposed, microbe-to-plant signal compounds, as the potential approach to address the challenges we
are facing. These compounds (e.g., lipo-chitooligosaccharide and thuricin 17) have been shown to increase
growth for diverse plant species, particularly when they are growing under stressful conditions, however, their
commercial development/utilization is far from complete. This review paper will expand the understanding of
using the signal interaction between crop and beneficial microorganisms not only to enhance plant growth but
also promote agricultural sustainability and a stronger bioeconomy.

1. Introduction
Global warming is by far the greatest threat mankind is currently
facing. According to recent data, an average increase in global temper­
ature of 2 ◦ C will lead to death of one hundred million people along with
extinction millions of faunal and floral species [1]. The Intergovernental
Panel of Climate Change (IPCC), in its fourth assessment report releasing
in 2007 and IPCC special report on climate change, desertification, land
degradation, sustainable land management, food security, and green­
house gas fluxes in terrestrial ecosystems released in 2019 expressed
that increased warming in last 50 years is, with a high degree of prob­
ability, due to intense emission of greenhouse gases (GHGs) including
carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) [2,3].
Carbon dioxide, among the pollutants, is considered to be the main
culprit due to the very high levels of anthropogenic emissions derived
from fossil fuel utilization [4]. Fossil fuels are being used to produce
energy for generating heat and electricity, and as transportation fuels,
contributing about 80% of GHG emissions produced globally [5]. The
use of fossil fuels is increasing along with the increase in human popu­
lation, which is projected to be around 9 billion by 2050 [6]. Thus, in
order to stop global warming, reduce GHG emissions and meet the en­
ergy requirements of modern civilization, fossil fuels need to be replaced
with renewable energy, which is a reasonable solution for global
warming and an efficient alternate source of energy [7]. Formation of
fossil fuels in the earth takes millions of years and their reserves are
finite, and subject to depletion through over exploitation [8].
Biomass, a naturally occurring non-fossil organic material containing
intrinsic chemical energy with potential to offset fossil fuel emissions,
could be a good alternative to fossil fuels [9]. Biomass resources from
agriculture, forestry and urban waste are comprised of a variety of

* Corresponding author.
E-mail address: donald.smith@mcgill.ca (D.L. Smith).
1
All authors contributed equally to this publication. With the exception of the corresponding author, the author order is alphabetical

https://doi.org/10.1016/j.rser.2020.110691
Received 29 April 2020; Received in revised form 20 December 2020; Accepted 29 December 2020
Available online 18 January 2021
1364-0321/© 2021 Elsevier Ltd. All rights reserved.
M. Antar et al.
distinct materials including wood, crop residues, sawdust, straw,
manure, paper waste, household wastes and wastewater [10].
Biomass-crop residues, if used as an alternate source of energy, have a
heating value of about 3 × 106 kcal Mg− 1; if compared with biofuels it is
about 50% that of coal and 33% that of diesel fuel and gives an estimated
fuel value of 18.6 × 109 J Mg− 1 (equivalent to 2 barrels (bbl.) of diesel)
[11]. As compared to fossil carbon commodities, bio-based products
release less GHGs, however the emissions vary with logistics, type of
feedstock material and technology used to produce end products. In
1970, the contribution of biomass as a source of renewable energy
supply received considerable attention worldwide due to the perceived
urgent need for sustainable energy self-sufficiency [11,12]. During
mid-1990s, in order to cope with global climate change and global
warming, there was renewed interest in biomass energy [13]. In this
millennium, we are investigating the subject of biomass as an alternate
and renewable source of energy largely for three reasons: i) to reduce
GHG emissions, in order to mitigate global warming and associated
climate change, ii) to minimize the overexploitation of finite reserves of
fossil fuels and iii) to reduce dependency on energy imports by gener­
ating sustainable renewable energy.
Over the next few decades, ongoing human population growth,
increased reliance on animal products and the associated elevated en­
ergy demand will require significant increases plant-derived bioenergy
and associated high-value products in the world markets [14,15], this
will reduce fossil fuel energy demand and associated greenhouse gas
emissions. The green revolution of the 1960s made a significant
contribution in enhancing plant production; more recently advance­
ments in biotechnology have, and will, help plants produce both more
and better biomass [14,16], due to improved plant efficiency to capture
light and convert it to biomass (carbon assimilation) with reduced fer­
tilizer and water inputs [17,18]. Therefore, it is crucial to investigate
more novel and lesser-known mechanisms for advancement in this area
[19]. These investigations and resulting understandings will help gov­
ernments and producers develop regulatory networks enhancing the
benefits of biomass production through a variety of plant species.
From an agricultural perspective, soil fertility now largely relies on
chemicals and physical properties in soils, origins and climate, rather
than biological activities in the soil [20]. To avoid this oversight, it is
necessary to deeply understand how chemical and physical properties in
soil are interrelated with biological activities, and all together are
affected by agronomic management practices. The understanding of soil
systems and their involvement in sustainable plant production could be
related to the interaction between microbe-microbe, plant-microbe, and
physical-chemical-biological [21]. Plant roots in the soil can serve as a
‘microbial hot spot’ due to their exudates and increasing nutritional
concentrations [22,23]. Microbe-to-plant signal compounds (e.g.,
lipo-chitooligosaccharides (LCOs) and thuricin 17) have been shown to
increase plant growth for a wide range of plant species, particularly
when plants are growing under stressful conditions [24–26]. The re­
ceptor for LCO is a LysM kinase for the legume-rhizobia symbioses; this
receptor system seems to have evolved for pathogen detection almost 2
billion years ago [27,28]. The microbe-to-plant signal in the N2-fixing
Frankia symbiosis remains to be identified but appears not to be an LCO
[29,30]. Thus, there are still more research need to be done realize the
potential of this very promising concept.
Due to the deterioration effects caused by the overuse of agriculture
land, the application of pesticides and herbicides, agricultural soils have
become relatively infertile, and thus result in low-productivity ecosys­
tems and, ground and surface water contamination, production of
greenhouse gases and soil acidification [31]. Therefore, solutions are
required that utilize strategies reducing the negative effects of interre­
lated variables without a loss in yield and sustainability. It has been
reported that the soil microorganisms have the potential to address
growers’ desires by increasing the crop efficiency in uptake capacity of
nutrients and water, leading to greater efficiencies of crop production
and, at the same time, greater yield [32,33]. As an approach with
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
substantial potential, plant growth promoting rhizobacteria (PGPR),
bacterial strains in soil and plant root systems, can enhance plant
development, growth and productivity [30,34]. These strains, depend­
ing on their niche, constitute three main categories including free-living
bacteria around the root zone (rhizosphere), root surface bacteria
(rhizoplane) and bacteria that live within plant roots (endophytic) [31].
However, environmental conditions in the soil and plant root system
may play important roles in allowing some bacterial strains to be present
in all three bacterial niche spaces [35,36]. Moreover, PGPR consortia are
recommended as biofertilizers for crops and it is an ideal alterative to
chemical fertilizer in that PGPR can help crops by increasing plant
productivity and providing protection against pathogens and other pest
organisms [37]. As synthetic fertilizers, which have a crucial role in
supplying almost half of the world population with food, despite several
diverse environmental conditions [38]. The demand for fertilizers is
increasing by about 1.9% per year [20], even though the expansion of
the use of fertilizer is leading to a series environmental problem
including eutrophication [39]. Therefore, urgent use of plant growth
promoting microbes is necessary to, among other things, facilitate the
availability of macro- and micronutrients to the plants, in order to
achieve significant biomass production improvement.
This review paper focusses on biomass utilization including bio­
energy, biofuel, bioproducts, and biochar. It also examines current
technological approaches utilizable to advance biomass production with
the focus on cropping system, genetic manipulation, phytomicrobiome
and its signalling compounds to produce sufficient biomass to sustain a
strong workable bioeconomy, reducing greenhouse gas emissions and
mitigating climate change progression.
2. Global biomass production
Biomass production and its potential to produce renewable bio­
energy varies among countries depending upon geography, availability
of resources, biodiversity, technology and economy. It is estimated that,
biomass could provide 3000 terrawatt hour (TWh) electricity by 2050
and could save 1.3 B t of CO2 equivalent emission per year, and for each
TWh energy produced 472.89 Kt CO2 is generated [40]. Globally,
renewable energy production increased to 18.6% by 2014. However, in
2018, renewable energy generation increased by 14.5%, slightly lower
than average growth (16%) of last decade. The Asia-Pacific region being
the leading contributor, accounting for 40% of global renewable energy
generation, followed by Europe and North America (Fig. 1, left). In
addition, China is among the countries that contributes the largest
amount of renewable energy with a production of 634.2 TWh followed
by the US and Germany, with renewable generation levels of 458.5 and
209.2 TWh, respectively. Moreover, the generation of renewable energy,
the consumption of renewables in 2018, worldwide, also indicated a
similar tendency as Fig. 1 (right) indicates.
During the past decade, renewable energy consumption patterns
have shifted drastically worldwide. China’s renewable energy con­
sumption has increased by 20-fold since 2008, however, much lower but
considerable increases have also occurred the US, Germany, Canada and
India (Fig. 2, left). Likewise, the increasing demand for biofuel, as an
alternate to conventional fuels, has eventually led to increased biofuel
production, worldwide. However, among the countries, the US con­
tributes the largest share of biofuel production, followed by Brazil and
Germany (Fig. 2, right).
However, increased biofuel production may affect agricultural
commodity (feedstock) prices. Globally, the major feedstock employed
for biofuel production are corn, wheat, sugarcane, soybean, rapeseed
and sunflower, which are directly or indirectly used in the food industry,
and currently, in international discussions is speculation that increased
utilization of feedstock for biofuel production will increase feedstock
prices and also food prices, due to increased demand and reduced supply
in the international market [42]. Food and fuel commodity prices, for
the first time since the green revolution, began to rise in 2001 and
M. Antar et al.
Fig. 1. Worldwide renewable energy generation by wind, solar and other renewables (left) and renewables consumption (right) in 2018 [41].
Fig. 2. Renewables consumption (left) and biofuel production (right) in 2008 and 2018 around the world [41].
peaked in 2008. During this period, the biofuel industry expanded
dramatically; ethanol (from corn and sugarcane) and biodiesel (from
soybean, oil palm and rapeseed) were produced two and six times,
respectively, that of 2001 levels [43], which was eventually recognized
as the main cause for the price increase of major feedstock (corn and
soybean) by twofold in the 2000s (the era of biofuel boom). In contrast,
the main reason for the price shock in agricultural commodity prices was
linked to crude oil prices. The continuous price increase of crude oil
from 2002 increased feedstock prices until the price drop after 2008,
when all commodity prices declined substantially [44]. Shrestha et al.
compared the food price index before and after the biofuel boom of the
2000s using real-world data [45]. Among the possible factors, they
found the highest correlation between the food price index and crude oil
prices, suggesting little evidence of food price increases due to biofuels.
Perhaps, food and oil are systematically connected, and their prices rise
and fall in tandem. The modern agricultural production system is highly
energy/fuel-dependent; mainly for operating farm machinery and
transporting inputs and outputs, and any increase in oil cost will even­
tually affect food prices, making them less affordable. However, there
are two strands of literature regarding food prices and biofuel produc­
tion: one that maintains oil market volatility is responsible for food
prices, while the other body of literature links increased biofuel pro­
duction to feedstock price shocks [46]. Biofuel demand and production
may have had a part to play in increased feedstock prices, but several
other factors also contributed to feedstock price rises, including global
economic growth, population growth, energy price inflation, exchange
rate fluctuations, adverse weather, trade policies, government in­
centives and feedstock production [47].
Despite these conditions, biofuels are often considered to be a solu­
tion to the energy crisis and an environmentally friendly approach to
sustainability; they have also been criticized predominantly in terms of
global food price increase, overexploitation and conversion of non-
agriculture lands into agricultural land, and diversion of food crops
and/or utilizing food crops for biofuel production [48]. It was suggested
that biofuel production and diversion of crops globally cause more
hunger, and this increase may less affect developed communities, who
spend less than 1/10th of their income on food, than the 2 billion poor
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
people worldwide who spend 50–70% of their earnings on food [49].
Biofuel production and utilization as a sustainable solution for energy
crises are crucial, but at what cost if it is a burden on food supply?
Certain things need to be considered and implemented to maintain an
appropriate equilibrium between bioenergy and food supply. Firstly,
food crop utilization as biofuel inputs should be reduced and/or new
techniques should be introduced to use and extract convertible sugar
from cellulosic non-grain parts of the crop to produce alcohol. For
example, corn is the main source of ethanol production, mainly from
corn kernels. But corn stalks and husks are considered wastes and are
rarely used for ethanol production; if ethanol is produced through
enzymatic hydrolysis of cellulose it can be prohibitively expensive for
commercial use due to high production costs. However, if appropriate
and inexpensive techniques are introduced to derive biofuels from
cellulosic feedstock, then it may increase the food supply and conse­
quently reduce food prices. In such scenarios, soil carbon sequestration
may be compromised; something crucial to be considered if imple­
menting such practices. Secondly, the utilization of food crops for bio­
fuel production should be reduced and/or replaced with cellulosic
material from non-food crops such as miscanthus and switchgrass,
however, overexploitation of lands and intense agricultural practices,
and mono-cropping may affect certain environmental aspects including
biodiversity. Thirdly, modern techniques such as genetics and growth
stimulating micro-biota could be employed as effective tools to increase
biomass production per capita. However, studies are being carried out to
practically implicate these tools in “biomass for biofuel” production, but
still, more relevant knowledge needs to be generated in order to increase
biomass production worldwide.
Recently, using biofuels derived from first-generation biomass
sources, often food materials, has been restricted by the European
Parliament resulting in a significant shift into the use of agricultural
residues to produce biofuel. At the same time, the European Parliament
designed and voted on draft legalisation to promote advanced biofuel
use and strongly encourage the market to use straw as a feedstock for
biofuel production [41,42].
Agricultural residues rank as the top source of biomass for sustain­
able energy production, due to their potential for minimal indirect land
M. Antar et al.
use change (ILUC) [43]. Research in Europe has indicated that bio­
ethanol production from straw has meaningful potential [44,45].
However, straw-derived bioethanol production and its sustainability
still need to be examined and developed to determine the effects of
bioethanol production on the agro-environment and overall greenhouse
gas emissions. In north America there is interest in recovering corn
stalks, with an emphasis on harvesting the upper parts of the stalks,
which are least recalcitrant to the bioconversion process, as they contain
less lignified material [46].
The European Union and the USA have set policies for renewable
energy. The aims of these policies are to increase shares of energy
generation by woody biomass and other sources of renewable energy.
The major regulatory policies, Renewable Energy Strategy (RES) direc­
tive in the EU and Renewable Portfolio Standards (RPS) in the USA,
provide more flexibility in meeting energy targets. Investigations of
renewable energy policies indicate that the effectiveness of existing
policies can be disrupted by other energy policies. Subsidizing low-
efficiency biofuel conversion systems or feedstock usage that only
cause marginal greenhouse gas emission reductions may not be the most
efficient use of public resources that could otherwise be used to develop
more efficient biomass conversion technologies. Renewable energy
policies, including woody biomass policies, need to be established
further and advanced approaches for environmental needs to be thor­
oughly reassessed [43].
Palm oil significantly contributes to the world’s vegetable oil pro­
duction leading to generation of large amounts of by-products and palm-
based biomass such as oil palm fronds, empty fruit bunches, palm
mesocarp fibre and palm kernel shells [50]. With regard to economic,
environmental and social aspects, value-added products including bio­
fuel, hybrid plywood, pellets and syngas are derived from palm-based
biomass through conversion technologies within a palm oil based bio­
refinery [48,49]. More recently, a new industrial system has been
developed to process oil-palm derived biomass and by-products in which
refinery, biorefinery and mill residues from palm oil, combined with
heat and power production, to introduce a new concept of palm oil
processing [44,51,52]. However, the major concern with palm oil is
sustainability. Environmental damage by forest burning is a major
problem; it is practiced when removing native forests for palm trees
plantation purposes for oil and biomass production. Historically, there is
a high degree of controversy regarding palm oil development, due to
sustainability concerns.
Globally, biodiesel and ethanol are becoming more attractive as
major alternative and ecofriendly transportation fuels, as compared to
diesel fuel and gasoline, respectively. However, their production costs,
and increasing prices over time are large challenges when competing
with the deep-rooted traditional oil market. Currently, despite higher oil
prices, renewable fuels (biodiesel and ethanol) are more expensive than
conventional fuels. The main reason, among a range of factors involved
in high biofuel prices, is the feedstock, which accounts for 75% of the
total product cost [51]. In addition, varying government policies and
incentives, processing technologies, land and labor costs in different
regions also contribute to biofuel price fluctuations [52]. Currently, the
prices of biodiesel are double those of conventional diesel, and ethanol
despite tax incentives, has not been able to compete economically with
gasoline. Brazil is the exception, in that it has ethanol prices near those
of gasoline. Although, biofuel prices vary among countries, depending
largely on the production of feedstock employed for biofuel production.
For example, ethanol production cost in US is much lower than Europe,
however, production costs in Brazil, using sugar cane-based ethanol, are
generally the lowest in the world. Developing countries with a warm
climate producing sugar cane-based ethanol, have lower production
costs than grain-based ethanol producing countries. Therefore, ethanol
produced in Brazil and India, with reduced production expenditures, is
becoming a cost-effective alternative to conventional fuels [53]. How­
ever, because of higher costs for renewable biodiesel and ethanol, may
not allow them to play a major role in global economic growth directly,
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
but they are well recognized for their impact on socioeconomic condi­
tions, sustainability, climate change, job opportunities, expanded agri­
cultural production, and predominantly, imports of and dependency on
fossil fuels.
2.1. Biomass potential in North America
2.1.1. Canada
Canada is rich in natural resources with enormous variety of geog­
raphy and landforms. Agriculture and the forest industry play a vital role
in countries with developing bioeconomies. Biomass from agricultural
and forest residues are being used by firms to produce bioproducts and
sustainable renewable energy. Currently, 27 million ton (Mt) of total
biomass produced, is being used by more than 200 Canadian firms
producing bioproducts making sales of $1.3 billion [54].
Annual agriculture residue production in Canada is estimated at
about 82.4 million dry Mg (excluding wheat hay and fodder corn) [55].
The major contributors to the country’s agricultural residues are Sas­
katchewan, Alberta, Ontario, Manitoba, and Quebec accounting for 96%
of Canadian crop production. Wheat straw is the dominant residue,
predominantly produced in western provinces and providing 92.6% of
total wheat straw supply, accounting for more than 40% of total agri­
cultural residues produced in the country. In addition, western prov­
inces account for a large proportion of residues produced from barley,
oat, canola, dry pea and flax. An overview of western Canada found that
37 Mt of cereal straw were produced annually. Although large volumes
were produced, there were significant requirements for soil conserva­
tion and livestock feeding. Requirements were 0.75–1.5 t ha− 1 for soil
protection against wind and water erosion [56]. In fact, after consid­
ering the requirements for soil conservation and livestock feeding, the
average residual amount available was only 15 Mt. However, there was
a major constraint to this large volume in that it had a large annual
variation with 2.3–27.6 Mt available annually [57].
Ontario and Quebec are the main producers of corn and soybean
providing 96.7 and 95.4% of total residues produced from corn stalks
and soybean straw, respectively [1]. The major constraints in Eastern
Canada are similar to western Canada, in that significant amounts of
material are required for soil conservation and that due to weather
fluctuations the ability to recover much of this biomass can be limited.
Canadian studies on crop residues suggest that a sustainable biomass
production will need to rely dedicated biomass crops to reduce risks of
biomass supply availability for commercial and industrial users.
Canada, with 347 M ha forest land, is the third largest forest area in
the world, with a great potential for biomass and bio-based products
[58]. Canada’s lignocellulosic feedstock from agricultural and forest
residues are estimated at 64–561 million dry t yr− 1 [59]. In 2007, the
contribution of biomass to domestic biofuel production accounted for 1
billion liter (B L) of ethanol and 97 million liter (M L) of biodiesel [60].
In 2015, 190 Canadian establishments involved in the production of
bioproducts, predominantly biofuels and bioenergy, accounted for 58.4
and 21.1% of total bioproducts produced. In 2015, the total sales from
bioproducts was estimated at $4.3 billion. Of these, biofuels, at 63.6% of
total sales, generated $2.7 B in revenue. In 2017, Canada’s forest
contributed roughly $24.6 billion to the country’s economy. The two
main forestry-derived bioenergy feedstock resources from forests are
harvesting residues and dead wood (standing dead timber) from natural
disturbance, predominantly insect attack [61,62]. Their future estimates
indicate an average annual production of 51 ± 17 and 20 ± 0.6 Tg yr− 1
from natural disturbances and harvesting residues, respectively [61].
In the Canadian context, there is a need to highlight and investigate
promising strategies to develop renewable energy sources, to displace
fossil fuels. For instance, the three important goals for development of
the biomass industry in northern BC include: building biomass energy
frameworks into existing networks that need access to natural gas,
providing a plentiful stock of biomass; extensively utilizing innovation
and planned approaches, and setting up sufficient funds to support these
M. Antar et al.
networks. The primary goal could be accomplished through open dis­
cussion and government backing. The second goal could arrive at
through associations among colleges/universities to building guiding
organizations. The final goal could be approached through a blend of
government awards and advance reimbursement programs, private
moneylenders, (for example, credit associations) and network reserves.
Providing a system of assets for biomass energy production, distribution,
and utilization in northern BC, and at other key areas inside Canada, will
be a significant step toward development of the considerable potential of
this energy source [41,45].
Canada depends intensely on its pipeline association with the US as
100% of Canadian natural gas exports are directed toward the US by
means of pipeline systems. Due to developments in local production in
the US, Canadian exports have been altered and this pattern, and this is
anticipated to proceed in the coming years. In effect, gas exports through
Canadian pipelines to the US dropped 7% between 2010 and 2014,
while gas imports to Canada via US pipelines expanded 21% during a
similar time span. Canada is presently assessing approaches to differ­
entiate their natural gas exports as confirming by the proposed Liquefied
Natural Gas (LNG) trade terminals in British Columbia province [53,54].
Given this situation, there are multiple considerations, including envi­
ronmental effects, growing energy demand, and the expected increase in
gas prices, forewarning us and indicating that we must change our en­
ergy supply systems. One of the best opportunities also lies in the
development and use of densified forms of solid biomass for heat and
power production. The EU market is emerging as a major consumer of
wood pellets for domestic, and commercial biomass heating systems and
for power generation. Raw materials for agri-pellets can be straw and
various other agricultural by-products. In addition to requiring more
specialized combustion systems, the storage and management ability of
by-products are important considerations in reducing the complications
of pellet production. Therefore, it is crucial to produce pellets that will
reduce energy input associated with transportation and improve com­
bustion parameters. Agri-pellet production is advancing slowly in Can­
ada even though it has a substantial raw material base. One reason for
this is that while we have various agri-pellet combustion furnaces and
boilers, these systems are relatively expensive and have had challenges
with functional reliability due to the high alkali content of densified
agricultural fuels. EU countries are helping with subsidies for the initial
investment to make the changeover to more advanced pellet heating
systems [41,45].
2.1.2. United States
The US is the largest producer of bioethanol, followed by Brazil.
Ethanol is an important pillar of the US fuel industry and the US econ­
omy. Ethanol (E10) is being used in all US gasoline, comprising 10% of
the total volume. Today, it is estimated around 56 B L are being used
annually in the total US gasoline pool. Corn starch is mainly used as a
raw material for bioethanol production [63]. In addition, corn is the
dominant crop cultivated across the country. In 2015, the area under
cultivation of corn is estimated around 38 million ha with a production
of 381 billion kg (1 bushel = 27.21 kg). Currently, 163 billion kg are
being used to produce 56 B L of ethanol. It is estimated that the corn
production and corn-based biofuel production will increase if ethanol in
the US, to the level of E20 (20% ethanol in the total gasoline pool) [64].
Ethanol production in the country is increasing substantially.
Comparing with ethanol production in 1998 (6.4 B L), US bioethanol
production has risen almost tenfold. In 2007, bioethanol production was
24.71 B L, which increased to 50.41 and 57.8 B L in 2010 and 2016,
respectively.
In the US, the biomass-based energy is contributing the larger share,
43% of total renewable energy produced [40]. However, cellulosic
biofuel is falling far short of production goals. In order to scale-up the
cellulosic biofuel utilization, the RFS (Renewable Fuel Standard) has set
ambitious goals for biofuel production. The RFS requires an increase in
biofuel use to 144 B L by 2022. Presently, there are about 200
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
biorefineries in the country with a potential of producing 60.64 B L
ethanol annually [65].
In 2018, the total renewable energy production in US was estimated
at 12,374,799 T J (British thermal units). Of which biomass-based bio­
energy contributed around 5,634,020 T J. Of these, biomass-derived
fuels including ethanol and biodiesel contributed 2397 T J, the
remaining being shared by wood derived fuels [38]. According to US
energy Information Administration (2019), the total corn and other
biomass inputs (feedstock) to produce fuel ethanol in 2018 was esti­
mated at 2,274,709 T J with a production of around 64 B L fuel ethanol
[66].
In addition to bioethanol, biodiesel is being produced in the country,
although not at the same level as ethanol [39]. In 2018, the estimated
biomass input including vegetable oils to produce biodiesel was esti­
mated around 253,214 T J. The total biodiesel produced was estimated
at 7.2 B L (US) [66,67].
Ethanol is mainly used in 10% blends with gasoline. However, it can
also be used in E85 form (15% gasoline and 85% of motor fuel blends)
where it can have a much greater impact on gasoline displacements.
Nevertheless, the automobile fuel market can be limited to grain-based
ethanol due to the availability of corn. For example, Corn grain can only
provide Midwestern states with 2/3 of ethanol required by regional
demands. This demonstrates the need for more advanced technologies
and additional resources, to expand bioenergy’s role in the trans­
portation sector [68].
2.2. Biomass potential in South America
2.2.1. Brazil
Renewable resources have a great contribution in Brazilian energy
matrix. In 2015, the share of renewable resources in the country’s total
energy matrix was 41.2%, of which sugarcane biomass constitute the
greatest portion, contributing 16.9%. The remaining component was
comprised of wood, charcoal and other renewable resources. Brazil, in
order to reduce its dependency on fossil fuels, focuses on generating
bioenergy from renewable resources. A major proportion of available
biomass is being used to generate electricity through thermoelectric
plants (stations). The main crop biomass contributors in the country are
sugarcane, corn, soybean and cassava. The total biomass produced from
these crops is estimated around 657.1 Mt yr− 1. Of these, sugarcane
production is superior in both production and bioenergy generation. The
production of sugarcane during the year 2008–2012 is estimated about
751.1 Mt yr− 1, providing 405.6 Mt of estimated biomass annually
(average) with an energy potential of 1,802,755 Gwh [69].
Brazil being the largest producer of sugarcane and ethanol produced
from it, has become a centre of attraction worldwide in terms of biofuel
production, predominantly ethanol. Sugarcane ethanol makes a signif­
icant contribution to the country’s bioeconomy. In 2013, the ethanol
production in the country was estimated around 28 B L and is expected
to increase substantially in the near future. Considering the importance
of sugarcane and biofuel produced from it, the Brazilian government is
focusing on the expansion of sugarcane and bioethanol production
throughout the country. Studies illustrated the impact of expanding
sugarcane and bioethanol production in 2030 could increase the na­
tional GDP by $2.6 B USD with an expected bioethanol production of 54
B L [70].
In 2017, sugarcane was cultivated on about 8.7 M ha with a pro­
duction of 633.3 Mt, providing 27.8 B L. In Brazil, sugarcane is mainly
produced in central and south regions. The major contributor of sugar­
cane and bioethanol in the country is Sao Paulo state, comprising 56.3%
of total Brazilian sugarcane production [71].
2.3. Biomass potential in Asia
2.3.1. China
Like other urbanized countries, China’s energy consumption is
M. Antar et al.
primarily fossil based, predominantly coal. The huge consumption of
coal for energy results in high CO2 emissions [72]. According to U.S
Energy Information Administration, among the energy consumers
worldwide China is top of the list. Therefore, the Chinese government
strives to limit GHG emissions by controlling the use of fossil fuels and
promoting renewable energy sources and diversifying non-fossil fuel
consumption. According to Canadian biomass magazine, China plans to
reduce coal consumption in rural regions by 50 Mt and produce 30 B m3
of biogas from crop residues and manure by 2030, which requires an
investment of CDN$ 7–10 B for building 3000–4000 facilities. China has
a very large potential for generating renewable energy from crop
biomass. Currently, China, through utilizing its renewable energy re­
sources, is the third largest bioethanol producer in the world. Since
2012, 1.5 Mt of bioethanol are being produced annually; the US and
Brazil are the leading producers of bioethanol [73].
Currently, China has 140 M ha of land area being used for crop
production. About 25% of the crop land is mainly used for paddy rice
production, the rest being used to grow crops like corn, wheat, tubers,
fibers and bean [74]. In 2010, China’s grain production was estimated at
about 570 Mt and straw production was 700 Mt. Excluding the amount
of straw used in industrial products, feed and fertilizers, 50% can be
made available to produce energy equivalent to 180 Mt of standard coal
[75]. Relevant studies have illustrated that every year around 500–800
Mt biomass is produced from crop residues with possibly 200 Mt being
available for biofuel production based on China’s projected straw pro­
duction during coming years by using average growth rate of grain
production from 1996 to 2013 [76]. The production in 2020, 2025 and
2030 will reach 841.00, 882.14 and 926.43 Mt, respectively.
In addition to crop residues, China’s forestry resources are increasing
in recent years. It is estimated that about 900 Mt of forest residues could
be available annually. Forest harvesting in China is estimated at about
379 M m3 annually, resulting in 239.53 and 139.47 M m3 of
merchantable and non-merchantable timber, respectively. Wood waste
and forest residues generated from timber processing in timber mills is
estimated to be around 201.1 M m3 and is projected to increase to 151.5
Mt by 2030. In 2016, only 45.7 Mt of biomass were used in all biomass
power plants [77].
Moreover, China is increasingly utilizing its marginal lands for bio­
fuel feedstock production by growing energy crops, due to limited
availability of cropland [78,79]. Investigations carried out by the Chi­
nese Ministry of Land and Resources (2003) and Ministry of Agriculture
(2007), indicated that the area of marginal land in energy crop pro­
duction was 23.72 M ha. It is presumed that these marginal lands will be
used to produce ethanol and biodiesel by growing energy crops like
cassava and Jatropha, respectively. In addition to these energy crops, it is
reported that switchgrass production in northern marginal lands is
estimated about 8–15 t ha− 1 [76].
2.3.2. India
India, the 7th largest country in the world, is a developing country
with a population of greater than 1 B and with meaningful potential for
bioenergy production [80]. Biomass production in India is estimated to
be 620–680 Mt annually. Of this, a small portion is used as animal feed,
household consumption and for small industrial units, however the
majority remains unutilized [79]. It is estimated around 100–140 Mt of
biomass is subjected to field burning, annually, equivalent to 1.5 Exa­
joule energy potential [81]. In agriculture dependent countries, field
burning of crop residues is frequent. According to the IPCC, 25% of
biomass, equivalent to 36 Exajoule of energy potential, is being wasted
in the world by field burning [82]. A recent study reported that in the
Indian state of Punjab annual welfare losses, in terms of health damage
by burning fields, amounts $1.1 M USD [80].
India being one of the largest fossil-based energy consumers and
GHG producers has introduced several incentive schemes throughout
the country, in order to reduce its dependency on fossil fuels by pro­
ducing bioenergy using unexploited non-fossil natural reserves
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(biomass). Currently, biomass contributes 32% of the country’s total
energy use [83]. It has been estimated that about 189 Mt of surplus
biomass from agriculture and forest residues is available every year for
electricity generation. Currently, the government of India is primarily
investing in renewable forms of electricity supply. According to the
Embassy of Denmark, India, about $9251 M USD is being invested in
biomass related projects generating 5000 M units of electricity. In
addition, Ministry of New and Renewable Energy plans to produce 10
GW of installed biomass power by 2022.
In India, the important biomass crops include Jatropha and sugar­
cane. Jatropha is an energy crop primarily used to produce biodiesel.
Jatropha is a stress-resilient tree: drought-resistant and grows easily on
degraded soils. It is predominantly cultivated in Asia, Africa and Latin
America. Among the various oil-bearing seeds, jatropha has a high oil
content (>37%) and the oil is of suitable quality, so that it has an
important place in biodiesel production. The oil from jatropha has low
acidity and viscosity, and higher stability than many other oil-bearing
seeds, making it more suitable for biodiesel production [84]. The most
common method for biodiesel production is transesterification of tri­
glycerides with methanol, in the presence of catalysts, to produce esters,
however, other methods are also employed for the purpose including
blending, microemulsion and pyrolysis [85]. The energy content of
biodiesel is lower than conventional fuel since it contains 10–11% ox­
ygen, hence it will require more fuel to generate the same energy as
conventional diesel. However, this chemical distinction makes it more
ecofriendly by reducing toxic emissions. For example, biodiesel B100
(pure biodiesel) reduces emissions of total unburned hydrocarbons,
carbon monoxide and sulphates by 67, 48 and 100% respectively [50].
Currently, the biofuel contribution to the country’s total fuel con­
sumption is very low, however, the Indian government is developing
plans around diversifying Jatropha cultivation to produce 20% of its
total diesel requirement by 2020 using biodiesel. To achieve the target,
around 11.2 M ha is planned to be rehabilitated by Jatropha plantation
[86].
India is the world’s 2nd largest producer of sugarcane, after Brazil.
During 2016–2017, sugarcane cultivation in India was estimated to be in
5 M ha, with a mean cane production of 70 t ha− 1. Sugarcane being the
primary source of bioethanol production in India, provides 22–25% of
ethanol t− 1 [87].
2.3.3. Malaysia
Malaysia is a south Asian country with significant potential for
biomass production. About 62% of the total land area is covered with
forest and approximately 4.9 M ha is occupied by agricultural produc­
tion, producing 168 Mt of biomass from oil palm (85.5%), municipal
solid waste (9.5%), the wood industry (3.7%), rice (0.7%) and sugarcane
(0.5%) [88,89]. In spite of this large biomass resource, 93% of total
energy consumption in Malaysia is dependent on fossil-fuels. It is pre­
dicted that the finite fossil fuel reserves in country will last for only
another 30–40 years. However, the Malaysian government has intro­
duced incentives for generating renewable energy resources, predomi­
nantly from oil palm as it is the biggest contributor of oil and biomass in
the country.
In Malaysia, oil palm plantation was estimated to be 4.3 M ha in 2007
providing 50–70 t biomass ha− 1. In 2005, biomass production from oil
palm was measured at around 55.73 Mt [28]. In 2006, Malaysia was the
2nd largest palm oil producer, following Indonesia, with a production of
15.8 and 15.9 Mt, respectively. Malaysia and Indonesia contribute 87%
of total global palm oil supply [5].
2.4. Biomass potential in Europe
Renewable energy plays a key role in both securing energy supply
and protecting the environment. Recognizing the need and importance
of biomass and its potential as an alternate and efficient source of energy
Europe has set a high and binding target of 20% as the share of energy
M. Antar et al.
production to be from renewable bioenergy by 2020 [89]. According to
a renewable energy progress report from the European commission, the
EU had reached 17.5% renewable energy by 2017, and so was closing on
the 20% by 2020 goal. Furthermore, the report emphasized a reduced
import of fossil fuel in 2015, resulting in a saving of €16 B. Following this
trend, this is projected to increase to €58 B by 2030. Currently, the
majority of EU countries are well on track to reach the binding union
target of 32% renewable energy in final energy consumption by 2030
[90,91].
Biomass is currently the largest source of bioenergy and fuel for heat,
electricity and transport in the EU. Prominent EU sources of biomass are
primarily from agricultural and forest residues. About half of the land
area of the EU is occupied with agriculture. According to a Science for
Policy report of the European Commission’s Joint Research Centre (JRC)
2018, the annual biomass production from agriculture, averaged for the
reference period 2006–2015, is about 956 Mt dry matter, including
primary products (grains, fruits, tubers) and crop residues with a pro­
portion of 54 and 46%, respectively. Residues from cereals including
wheat (149 Mt yr− 1), maize (80 Mt yr− 1), rapeseed (54 Mt yr− 1) and
barley (50 Mt yr− 1 contribute 74% of total residual biomass produced.
Among the member states of the EU, France and Germany are the largest
producers of residues, at 84 and 60 Mt yr− 1, respectively, and together
with Poland, Italy, Spain, UK and Romania make up about 75% of total
crop residues produced [91,92].
In 2016, the total biofuel consumption in EU was estimated at 13,840
ktoe (Kilotons of Oil Equivalent). Out of which biodiesel contributed
80% of the total bioenergy consumption with an energy potential of
11,083 ktoe, followed by bioethanol with a share of 19% in sustainable
consumption of biofuel. Of the total biodiesel consumed in the EU, 74%
is from the continent’s feedstock material, mainly from rapeseed
(~38%), and the remaining 36% is biodiesel from palm oil imported
from Indonesia and Malaysia. Rapeseed imports from Australia, Ukraine
and Canada constitute 11% of total biodiesel produced in the EU [91,
93].
In addition, EU feedstocks provide 65% of total bioethanol
consumed. Of these, 64% is largely from agricultural crops such as
wheat (~25%), corn (22%) and sugar beet (17%). Canada is third
among the major providers of feedstock (1.6%) for bioethanol produc­
tion in the EU [90].
In addition to agricultural biomass, European forests play a key role
in the continent’s bioeconomy by providing crucial resources for
biomass supply. The role of forests, in terms of future climate security
and bioeconomy, by provision of raw material, is quite considerable.
Globally, annual wood production from forests is estimated at 3.7 B m3.
The area under forest land in the EU is estimated to be 161 M ha
[FAOSTAT, 2018]. Biomass from EU forests is mainly from fuel wood
and residues from logging and wood-processing, and these are the major
sources for generating bioenergy.
In 2016, the woody biomass supplies from forests and other wooded
land contributed about 60% (82 Mtoe) of total biomass used for energy
purposes. The woody biomass potential and availability in majority of
European countries has been assessed. The results illustrated that the
annual total biomass production from European forests is about
357–551 Tg dry matter [94,95]. In 2016, the global production of wood
pellets reached 29 Mt, and of these more than 50% was produced in the
EU [96]. According to a recent IEA report on bioenergy (2017), the
overall wood pellet production worldwide is estimated to be 26 Mt with
an increase of 14% per year since 2010. Considering the increased trade
of wood pellets worldwide and its global demand, the EU is the largest
producer (54%, 2014), followed by North America; Canada and the US
(35%). The remaining 11% share of global wood pellet production is
mainly from Asia, the Russian Federation, Australia and Latin America
[97].
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
3. Biomass utilization
The world’s energy markets have a heavily reliant on coal, oil and
natural gas, collectively known as fossil fuels. By-products of fossil fuel
burning include a variety of toxic air pollutants and CO2, thereby posing
a major threat to the human’s health and wellbeing and contributing
significantly to global warming and environmental degradation. Once
coal mining started, in the 18th century, coal became the main fuel of
the industrial revolution; previously, the human need for energy was
totally met by wood and charcoal which are biomass [98]. These
constitute only known organic, renewable carbon resource that is suf­
ficiently large to act as a substitute for fossil fuels is biomass [99].
Biomass energy is a type of renewable energy and, as opposed to fossil
fuels, it can be used directly or after conversion to other forms, releasing
the amount of CO2 that the biomass recently captured from the atmo­
sphere during its growth. That is why biomass is considered ‘carbon-­
neutral’, which has attracted worldwide interest in using biomass as an
alternative for fossil fuel derived energy sources, particularly as a source
of liquid fuels and chemicals (methanol, ethanol, biodiesel, etc.) [98]. As
of now, a wide range of sectors benefit from biomass: either as food/feed
to generate electricity, heat and fuels or as a material and resources in
the industrial fields mainly in the wood, paper and chemical industry.
3.1. Bioenergy
The current pattern in the energy sector is economically, environ­
mentally and socially unsustainable. Greenhouse gas emissions in the
world have increased twofold since 1970 [100] and enhanced oil de­
mand has resulted in concerns regarding the security of supplies, indi­
cating a situation where an urgent decision is needed. One potential
approach to address this issue is bioenergy. Bioenergy is the oldest and
largest source of renewable energy; it has generally been entirely
derived from waste materials including lignocellulosic residues from
forestry (logging, thinning, and processing residues), agricultural (har­
vest and processing residues), and food and municipal solid wastes.
Furthermore, bioenergy-grown crops, virgin lignocellulosic biomass and
algae are used as raw materials for production of bioenergy [101];
nowadays it can provide heat, power and transportation fuels, ac­
counting for approximately 10% of world total fundamental energy
supply [102] and this proportion is increasing by about 2.5% annually
around the world [103]. The amount of the bioenergy released from
bioresources entirely relies on the type of feedstock: woody biomass
with higher lignin content (23.9–32.0%) is more solid and denser
(350–490 kg/m3) than agricultural biomass (108–130 kg/m3), which in
itself makes it more resistant to microbial and enzymatic actions than
agricultural biomass (Zhu and Pan, 2010). For bioenergy production,
specifically biofuel, a higher level of cellulose in biomass is required to
produce more volatiles [104], which is necessary for ignition and
oxidation during combustion [105] whereas lower hemicellulose and
lignin content is an appropriate feature of feedstock since lignin acts as a
chemical glue, making it difficult to separate cellulose from hemicellu­
lose, and prevent the production of amorphous cellulose from crystalline
cellulose. Moreover, lignin inhibits access of cellulase to cellulose,
causing inhibitory effects on biological conversion of biomass [106].
The most common bioenergy use is primary heat and cooking in less
economically-developed countries [107]. The main issue related to this
sort of bioenergy is that the biomass is sourced unsustainably, resulting
in forest degradation. Moreover, the efficiency of burning biomass in
this way is between 10 and 20%, with considerable indoor pollution
[108], whereas large-scale combustion plants produce heat efficiently
and in a way that is competitive with fossil fuels. Commercial systems
for generating heat from biomass range up to extremely large boilers in
the 10 MW (megawatts) range, which are mostly employed within in­
dustry, and small ones that supply heat for household consumption from
timbers, wood chips or wood pellet feedstock. Biomass can be also
transformed into electric power through a variety of methods but most
M. Antar et al. Renewable and Sustainable Energy Reviews 139 (2021) 110691

biopower plants utilize direct combustion techniques, burning biomass Table 1

specifically to deliver high-pressure steam that runs a turbine generator Comparison the potential of different biomass sources for biofuel.
to create power [89]. The electric output of a steam turbine relies on its Biomass Feedstock Specific characteristic References
capacity and commonly, electric turnover is between of 15–35% [109]. category examples
The amount of CO2 for bioelectricity generation depends on how Pine • Less ash content than crop [144,145]
effective the conversion technology is and how much fossil fuel is residue
consumed to produce biomass; good agricultural and forestry manage­ Cedar • High density of the biomass
ment is a key factor. In one study, cost of production and carbon making more economical
transportation
abatement per unit of generated electricity for several feedstocks was Forest biomass Poplar • Flexible harvesting time
estimated in the US, which is significantly lower than fossil fuels; cotton and waste
stalk is the most GHG intensive (325 kg CO2e MWh− 1) preceded by corn Oak • Various type of biomass based
stoves (235 kg CO2e MWh− 1) whereas pine chips emitted the least GHG on topography, management,
climate and disturbances
(134 kg CO2e MWh− 1) for a similar amount of electricity [110]. World
Willow
electricity demand will burgeon from roughly 20,000 TWh (tera Mouha
watt-hour) in 2009 to 42,000 TWh in 2050, due to the growing sus­ flowers and
tainable energy economies and it is estimated that the share of renew­ seeds
able power will increase from 19% in 2009 to nearly 60% in 2050 [111]. Mahula
Switchgrass • High biomass production [146–148]
Miscanthus • High cellulose content (25%–
3.1.1. Biofuels 40%)
There is an expanding need for energy and related infrastructure to Perennial grass Reed canary • Less dependency on fertilizers
fulfill human social and financial advancement, welfare and wellbeing. and pesticides
Giant reed • Quick and easy growth
Returning to renewable sources to assist in relieving climate change
Indian grass • Having potential ethanol
challenges is an excellent approach that should be an achievable way to yield 160–450 L/ton of
meet the energy demands of future generations eneration’s energy de­ biomass
mands [112]. Fuels from biological sources (Table 1) such as grains, • Carbon sequestration
sugar crops, oil crops, starch, cellulosic materials (grasses and trees) and • Improve soil quality
• Growth in marginal lands and
organic waste defined as biofuels [113] which could be in three main
abandoned areas
classes: solid, liquid and gaseous phases: (i.) firewood, wood chips, wood Corn stover • Does not compete with food [149–151]
pellets and charcoal are types of solid biofuels, used since the dawn of and stalk production
human culture; wood and other plant substances were used for heating Sugarcane • Reduced dependency on
Bagasse forest biomass, leading to less
and cooking [126] although those materials are now largely burned in
deforestation
developing countries. (ii.) Biogas is another type of biofuel derived from Agricultural Sorghum • Short harvest time and easy
anaerobic digestion of biomass, a renewable gaseous fuel with potential residues straw availability
to replace natural gas [114,115]. Considering the available agricultural Wheat straw • Comparable cellulose content
feedstock and domestic wastes, it is estimated that annually more than (30%–45%)
Rice straw
1000 B Nm3 of biomethane could be released to produce electricity and
Red algae • High photosynthetic [152–154]
heat [116]. Switchgrass is a tall warm season perennial grass with po­ efficiency
tential to produce more biomethane than other crops through anaerobic Aquatic plants Green algae • Short life cycle
digestion [117] which is an environmentally-friendly, low-cost method Brown algae • Low nutritional requirement
• High potential of biomass
in which microorganisms convert raw biomass to methane and carbon
yield
dioxide [118]. However, the decomposition of its lignin only by mi­ • No competition with
crobes does not lead to full production of biogas, hence, chemical, mi­ terrestrial plants
crobial, mechanical or thermal pre-treatments are carried out [119]. • Have bioremediation
Methane output for most grasses was enhanced with delayed harvest capability
Food waste • Mutual relation between [155,156]
whereas the effect of harvesting time in cool and humid conditions, like
waste management and
Eastern Canada, is different. The maximum methane yield is derived energy security
from switchgrass harvested the in mid-summer or in early-fall; more Municipal Residences • Approximately 21 million
methane, approximately 25% more, could be obtained with a two-cut solid waste Leftovers ton/year for an urban area
with 200 million of
strategy in comparison to harvesting once in late summer [120]. Syn­
population
gas is another type of gaseous biofuel manufactured by gasification or Garden waste • Potential of 154L/ton of
pyrolysis of feedstock [1]. After purification, syngas is used as a raw biomass
material for synthesizing fuel for transportation, methanol, ethanol,
methane, dimethyl ether and other products [121]. (iii.) Biodiesel, uti­
lized in place of diesel, and bioethanol, as a substitute for petrol/gaso­ be considered for biomass production, if applied. Therefore, a life cycle
line, are the two most frequently used forms of liquid biofuels [122]. analysis (LCA) including the assessment of raw materials, land use
According to the latest U.S. Energy Information Administration (EIA) change, biofuel manufacturing and final use for each type of biofuel is a
projections, the utilization of global petroleum and other liquid fuels necessary tool to determine the efficiency of biofuel over fossil fuels
will rise by 38% between 2014 and 2040 [66,67] and the transportation [124]. As an example, the LCA of soy biodiesel in US represented a
and industrial sectors will consume 92% of the worldwide liquid gaso­ reduction of fossil energy consumption by 80% and a 66–72% decrease
line in 2040 [123]. This burgeoning demand drives the search for a in total GHG emissions, compared to fossil diesel. The farming process
range of biomass feedstocks for production of fuels which possess the plays a key role in calculation of GHG emission of biofuels, for instance,
ability to replace fossil fuels. The combustion of biofuel in engines re­ canola biodiesel is produces considerably more GHG, relative to soy
sults in CO2 emissions which are offset by the photosynthesis of the biodiesel for biomass production, which is 18.5 g CO2e/MJ in compar­
plants from which it was derived. However, GHG emissions can still be ison to 9.2 g CO2e/MJ for soybean. This is mainly because of nitrogen
significantly high since fossil fuels are used for agricultural machinery fertilizer application because canola cannot fix atmospheric nitrogen,
and transportation. In addition, fertilizer, pesticides and herbicides must while soybean can [125]. First and second-generation biofuels are

8
M. Antar et al.
successfully deployed by industry while the third generation is still
relatively new, but has a promising future [126]; biofuels beyond first
generation are often collectively referred to as advanced biofuels. The
first generation of biofuels, such as ethanol are prominent and are
created from food crops based on sugar, starch, or plant oil such as
sugarcane, corn and beet, wheat or soybean (oil) in Brazil, the United
States of America (USA) and Europe, and oilseed rape in France and
Germany and palm oil in Asia and Africa. Most of the commercially
created biofuels are derived from these feedstocks whereas conversion
of lignocellulosic materials such as cellulose, hemicellulose, and lignin
produces second generation, or advanced, of biofuels [127,128]. Third
generation biofuels are based on algae and marine plants (seaweeds).
The algae-produced biofuels have tremendous potential to be a substi­
tute for fossil fuels as they can use sewage, wastewater and saltwater,
growing in areas inappropriate for crops produced for first and
second-generation biofuels [129]. Nonetheless, the operational steps are
costly and still require research and development to make a third gen­
eration of clean energy sufficiently efficient. This type of feedstock is
able to grow three or four times faster than terrestrial plants and pos­
sesses relatively equivalent cellulose contents to agriculture or forest
residues while it has a low lignin and hemicellulose contents [130]. The
untreated macroalgae has approximately 76% cellulose conversion
whereas treating feedstock with peracetic acid followed by ionic liquids
causes almost complete cellulose conversion [105]. If the methods of
turning biomass waste into useful biofuels do not require excessive
levels of energy, biofuel production could be significantly advantageous
over crude oil and coal. There are several available technologies and
methods for biomass energy conversion. Two common methodologies
for converting biomass to fuel are thermochemical and biochemical
transformation, plus transesterification is used for production of
first-generation biodiesel [131]. The first technique utilizes heating for
biomass decomposition [132,133], while the latter utilizes microor­
ganism or enzymes to transform biomass to biofuels [134–136]. In brief,
the general process of biofuel production includes two main steps; (i.)
decrease the oxygen of the feedstock to enhance energy density and (ii.)
formation of C–C bonds between intermediates derived from biomass to
increase the molecular weight of the final product [137]. The world’s
biofuel production has increased sharply from 9.2 Mt of oil equivalent in
2000 to 95.4 Mt of oil equivalent in 2018 [138–140]. This thriving
growth has been triggered by policies that encourage the utilization and
production of biofuels due to the perception that it might offer energy
security and scale back greenhouse gas emissions in relevant sectors.
3.1.2. Biofuel through the Canadian lens
Canadian forests, agricultural land, and municipal wastes contribute
58, 39 and 3%, respectively, of predicted total energy content (5.3 EJ
yr− 1) from biomass [133,140–142]. It should be noted that residues
from harvesting of forests, agricultural fields and municipal waste are
still largely untouched [141]. Thus, Canada’s biofuel sector isn’t feed­
stock limited. Almost 176,000 ML of biofuels can be generated sus­
tainably in Canada from all yearly produced biomass. If only 10% of
total biomass from harvest residue of forests, agriculture and municipal
wastes is utilized for biofuel, approximately 17.6 ML of biofuels could be
produced, whereas the current production is around 2.0 M L [142]. The
biomass assets could contribute more to diminishing GHG emissions.
According to data released by provincial and federal government sour­
ces, ethanol utilization has expanded from about 1.7 B L in 2010 to 3047
M L in 2017, and biodiesel consumption has increased from approxi­
mately 123 M L in 2010 to 376 M L in 2017, resulting in 34.3 Mt less CO2
emission between 2010 and 2017 [143].
3.2. Biochemicals/bioproducts
Other than production of bioenergy and biofuels, biomass has been
employed for production of specific biomass-derived chemicals (bio­
products). It is estimated that near 50 Mt of bio-based and polymer
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
materials are currently produced around the world [157] indicating the
potential for economic sustainability [158] and reduced dependency on
fossil fuels, causing meaningful reductions in GHG emissions. Oil and
gas, nonetheless, are still broadly exploited for organic chemicals and
polymer manufacturing, which is not in keeping with the current
growing movement toward reduced pollution, environmental degrada­
tion and resource depletion. A wide diversity of bio-chemicals including
products with long records of bio-based history such as citric acid, ma­
terials newly introduced to the markets such as propylene glycol, and
those with significant growth and development obtained from biomass
[159]. The majority of chemicals are derived by specific building blocks;
interestingly, all those blocks could be replaced by their counterparts
from biomass known as bio-derived building blocks such as formic acid,
lactic acid, glycol, propylene, succinic acid, furfural, xylitol, sorbitol,
lysine and ethylene glycol. In step with the concept of tree products in
the chemical industry, a single chemical precursor can be converted to a
number of derivatives, linking chemicals along the value chain [160].
Therefore, a broad range of value-added materials or chemicals from
biomass could be produced either as final products such as adhesives,
lubricants, pest control agents, paints and coatings, pigments and inks or
intermediates for further processing such as manufacturing of bio­
plastics [161,162], bio-composites [163,164] and ingredients for cos­
metics, fragrances, food, pharmaceuticals and medical devices.
3.3. Biochar
Biomass with definite chemical structure can be used in numerous
conversion processes such as pyrolysis to manufacture multifunctional
products such as biochar [165]. The International Biochar Initiative
(IBI) has described biochar as a carbonizing material with a high carbon
content that is degradation resistance and derived from
thermo-chemical decomposition of bio-based feedstock by pyrolysis or
gasification in absence of added oxygen [166]. Biochar is widely
applicable in industry, agriculture and the natural environment. It can
be applied as a soil supplement, to forage and silage crops, and as a
water treatment [167,168]. Biochar has been seen as a potentially suc­
cessful approach to reducing CO2 concentration, since it retards return
of soil fixed carbon to the atmosphere. Biochar also hinders the emis­
sions of other greenhouse gases such as N2O and CH4 from soil
[169–171]. This is maybe due to the fact that biochar increases the
adsorption and confinement of ammonium in soils causing reduction of
the available amount of nitrogen for denitrification [172,173]. This soil
amendment can immobilize contaminants in the soil and sewage as it
provides binding sites for pollutants because of its high absorption rate
and resistance to microbial degradation [174,175]. Biochar is currently
being marketed as a way of launching a “doubly green revolution” [176]
by tackling GHG emissions from soil organic matter and sustaining food
security at the same time [177–180]. It also has positive effects on
physical aspects of soil such as density, porosity, structure and texture
[181–183], and also chemical features including cation exchange ca­
pacity (CEC), pH, soil carbon, nutrient cycling and water retention
[184–187]. Furthermore, application of biochar causes a range of effects
on soil biota by providing a carbon substrate [188], release or absorp­
tion of substances which benefit [189] or hamper microorganisms
[190], or extending microorganism niche spaces through providing
habitat for them [176,191]. Thus, biochar has potential as a valuable
tool on farms leading to sustainable agriculture particularly in tropical
regions [192,193] where there is a lack of soil organic matter because
climate conditions causes rapid degradation [194]. For instance,
metal-analysis of data collected from 16 fields and pots reported in­
creases up to 30% in yield when biochar amendments enhanced soil pH
by 2.0 in acidic, medium- and coarse-textured soils [195]. Likewise,
biochar soil amendments increased corn yield due to enhanced soil
water retention, nutrient availability and modified root system devel­
opment [196–198]. In addition to tropical sites, there are reasons to
think that biochar might be beneficial for temperate areas. Utilization of
M. Antar et al.
biochar in soil provides N for plants by higher N retention, reducing
leaching of plant available NH+ 4 and NO3 which is particularly impor­
−
tant for non-nitrogen fixing crops, and modifying the N cycle due to its
effects on soil microorganisms. For instance, after biochar application
rice yield was 10–20% greater than control plots, in the first and second
cropping cycles, in China because of the increased soil pH, soil organic
carbon, and total nitrogen and improvements in soil bulk density [199].
Similarly, the response of corn, soybean and switchgrass to pine wood
biochar at rates of 0, 10, and 20 Mg ha− 1 were studied in loamy sand and
sandy clay loam soils in southern Quebec. On the loamy sand, applica­
tion of 20 Mg biochar ha− 1 improved corn yields by 14.2% compared to
the control; this effect was not present on the sandy clay loam soil.
Biochar did not influence nutrient availability or yield for soybean or
switchgrass plots on either soil type. In line with these results, impacts of
biochar utilization are dependant on soil texture and crop type [200].
Apart from agricultural benefits, biochar can be used as ancillary ma­
terial in methane fermentation and composting [201]. Moreover, bio­
char has been implicated as a substrate in hydrogen formation, as a filter
in pyrolysis and gasification or as a fuel when pelletize [97,202,203].
4. Approaches for attaining improved biomass production
Some of the major challenges we are facing in the 21st century relate
to the need to increase food production while decreasing inputs, to
decrease greenhouse gas emissions and their effects on the environment
and to adapt to the climate change that has occurred and will occur
[204]. Addressing these problems is a long-term and complicated proj­
ect, but it is not an impossible challenge; there are advanced technolo­
gies being applied at various locations around the world. Whereas,
modern technologies, could reduce breeding time and allow for rapid
selection of novel crop varieties or species with exceptional
performance.
4.1. Cropping system
The cropping system deployed is as important as the selection of
high-performance genotypes. Rotation systems have dramatic effects on
soil architecture, utilization of soil resources and even the over all agro-
ecosystem. For example, intercropping systems play an important role in
the soil fertility, crop diversity and improvement of biomass yield
through planting two or more crops at the same time on the same land
surface [205]. It is also feasible to grow a perennial crop as part of an
intercropping system, instead of conventional cultivation through
growing annual crops; this could be particularly effective on marginal
lands, because this could decrease the input of nitrogen fertilizer while
contributing to the enrichment of organic carbon in soil [206] and also
enhancing soil biodiversity [207]. Thus, cropping system can have
positive effects on the N utilization and biomass yield improvement, as
when Sida hermaphrodita was intercropped with Trifolium pretense [206].
Moreover, the cover crop rotation can also achieve the goal of biomass
production improvement through combining winter cover crops with C4
and/or C3 crops [208], which improve water and nitrogen use
efficiencies. This approach can also result in biomass production on the
land surface during the off-season gap, resulting in more efficient use
sources and leaving soils less vulnerable to erosion.
Planting patterns affect crop root growth, which is an important
factor in nitrogen (NH+ 4 , NO3 ) uptake [209]. Improvement of nitrogen
−
use efficiency can be most effective through use of biological N fixation,
a critical N source in more sustainable agricultural systems [210]. It has
been reported that the cereal crops take up more N in legume-cereal
intercropping systems. For instance, in legume-cereal intercropping
systems, nitrogen use efficiency is increased by 1/3 and there is a 30%
grain yield enhancement, when compared with a monoculture system
[211], which could be explained by nitrogen fixation of legumes causing
higher nitrogen use efficiency. In contrast, increasing use of synthetic N
fertilizers significantly decreased nitrogen use efficiency. Therefore,
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
selecting an appropriate cropping system could decrease N fertilization
inputs and improve the nitrogen use efficiency, achieving better eco­
nomic value [212] and also reducing emissions of the potent greenhouse
N2O.
4.2. Technologies for improving biomass production
It is important to fill the yield gap while also developing methods
that allow lower inputs with less environmental damage. This aids in
achieving goals such as improved water and nitrogen utilization. Ni­
trogen use efficiency is considered an important indicator for crop
production sustainability and environmental impact, because high ni­
trogen use efficiency can enhance crop biomass production with lower N
application levels, while also decreasing the pollution risk [213],
including production of the potent greenhouse gas N2O. However, ni­
trogen use efficiency does not increase with the N input, which also does
not contribute to crop production as much as one might expect/hope.
One of the main reasons is that over-using N fertilizer diminishes the
stability of agricultural systems and has detrimental impacts on the
environment.
In addition to the effect of cropping system on nitrogen use effi­
ciency, the presence of soil fauna (protozoa, earthworms) and micro­
organisms (fungi and bacteria) in soil have influences on the physiology
and overall biology of plants [214]. During the plant growth period,
microorganisms colonize the rhizosphere and communicate with roots
to improve soil fertility by metabolizing the N that is not absorbed by
plants [215]. NO−3 and NH+ 4 uptake could be enhanced by interactions
between plants roots and some microorganisms such as N2 fixers, to
improve N availability, arbuscular mycorrhizal fungi (AMF) which play
a key role in P uptake [216]. For instance, the extraradical mycelium
produced by Rhizophagus intraradices, an arbuscular mycorrhizal fungus
species, can stimulate NH+ 4 absorption by plants, from soil, to improve
the nitrogen use efficiency [217]. In addition, PGPR are also able to
impact nitrogen use efficiency through enhancing N availability in the
rhizosphere and affecting N metabolism in plants, which both respond to
the crop yield [218]. For example, inoculating Achromobacter strain
U80417 onto roots of oil seed rape (Brassica napus L.) improved the
transport levels of NO−3 /NO−2 inside plants [219], providing key evi­
dence that some PGPR are able to improve nitrogen use efficiency and
contribute to the enhancement of biomass production. It has also been
found [220] that inoculation with N2-fixing bacteria can enhance wheat
yield through providing more nitrogen to wheat plants.
Furthermore, microorganisms affect survival of pathogenic bacteria
in soil, and prime plant defenses against biotic and abiotic stresses [24,
221]. Plant-associated micro-organisms, such as strains of Bacillus and
Pseudomonas, could enhance plant growth under stressful environments
by producing stress-related plant growth hormones, such as IAA, and
exhibit metal resistance ability and may reduce metal toxicity and ef­
fects of metal translocation within plants via secretion of acids, proteins,
and other chemicals [222,223] and through enhancing ACC deaminase
activity [224]. In addition, AMF are also involved in plant interactions
with soil toxic metals, and improving uptake of nutrients such as P, and
other relatively immobile soil nutrients such as Cu and Zn [225,226].
Moreover, as the published data indicate, the ability of arbuscular
mycorrhizae is affected by environmental conditions, fugal species and
plant characteristics, when the plant is experiencing an abiotic challenge
such as metal stress [225,227].
Although microbes have significant effects on the N cycle in soil-
plant ecosystems, cultivation or any disturbance of soil impacts N
transportation and assimilation through changing the microbial com­
munity’s diversity and components [228]. The use of crop cultivation
system, such as intercropping or cover crop, can be a feasible and
valuable strategy, associated with appropriate plants able to fix nitrogen
and utilize AMF symbiotic associations, which are also promising agri­
cultural technologies, to adequately utilize the soil resources and
decrease fertilizer nitrogen input. For instance, applying leguminous
M. Antar et al.
plants as one of the components of a cover crop could be an efficient
approach to improving nitrogen use efficiency through the beneficial
effects on AMF and PGPR, and their effects on soil N availability and N2
fixation (Fig. 3). In addition, cover-crop cultivation can improve the
ability of mycorrhizae to colonize plants and, again, improve nitrogen
use efficiency, leading to enhanced crop yield [228]. Thus, the combi­
nation of better microorganism activity and optimum agricultural
practices also can be used as a promising strategy to take full advantage
of available nitrogen to achieve substantial increases in crop production.
Currently, the improvement of microorganism activity and
enhancement of nitrate signaling could be the most effective strategy to
improve biomass production, though the combination of appropriate
cropping system components [229]. Specifically, it is well known that
diversity of plant colonization within cover and/or intercropping sys­
tems with AM fungi significantly increases potential yield. Related ap­
proaches can be promising alternatives for increasing plant production,
based on the same amount of N fertilizer supplement. Furthermore,
inoculation of AMF enhances the coding and expression of nitrate
transporters. For example, inoculation of Phyllobacterium brassicacearum
STM196 can stimulate NRT2.5 and NRT2.6 expression in Arabidopsis
plants [230]. Another new method to increase nitrogen use efficiency is
to regulate related gene expression. Increases of 50–60% in crop yield
are reported to be from genetic improvement of crops such as maize (Zea
mays L.) [231,232]. Thus, improvement of nitrogen use efficiency en­
hances crop yield and uses land and N efficiently by using of appropriate
cropping system, breeding approach, manipulating the expression of key
genes related in N uptake, assimilation and transport.
4.3. Modern genetic crops in relation to biomass production enhancement
Biomass production improvement depends not only on the selection
of crops that perform well in the area where they will be produced and
under advanced agricultural practices, but also needs to be based on the
prevailing environmental conditions. Currently, climate change is un­
predictable, and the crops of interest will be dealing with increased
levels and frequencies of both abiotic and biotic stresses. Therefore, the
development of novel crops, or novel forms of existing crops, both in
Fig. 3. Examples of the effect of PGPR and AMF on crop production in cover cropping systems, simplified from Coskun., et al. [228]. The root exudates produced by
specific plants trigger the response of specific beneficial microbes which send return signals to plants and improve plant growth; this process constitutes a symbiosis
between microbes and plants. In the meantime, the N2-fixation by microbes supplies extra N to the plant without the ability to self-supply the nitrogen.
11
Renewable and Sustainable Energy Reviews 139 (2021) 110691
terms of inputs and composition is critical to adapting to the changing
environments. Modern genetic engineering techniques are a critical
approach to solving this problem at the bioeconomy level [248].
4.3.1. Genetically modified crops for biomass production improvement
The use of genetic modification (GM) biotechnology involves
modifying the genes in an organism and/or how they are expressed, such
that the organism can then carry out specific metabolic activities giving
the organism enhanced utility [233,234]. The transfer of a specific gene
element or entire gene with yield or quality enhancing activity results in
an elite GM variety [235]. Microbes and microbial enzymes better able
to process biomass can and have been developed, and can be applied to
GM biomass energy crops to reduce energy and economic costs associ­
ated with biomass production and processing; these approaches can also
be used to generate biomass crops with high value attributes such as
resistance to insect pests [236]. Advances in GM biotechnology have
considerable potential in providing an key underpinning to the devel­
oping bioeconomy [236,237]. Currently, GM technology has been
extensively utilized in development of food crops with higher yields,
better pest (insect and disease) resistance and enhanced stress (biotic
and abiotic) resistance [236]. Globally, total food production increased
on the order 370 M t between 1996 and 2013 because of GM technology
utilization in crops such as corn, soybean and canola [234]. During the
same period, GM crops also resulted in substantial environmental ben­
efits through reductions in the application of both herbicides and pes­
ticides, resulting in estimated 37 and 18% reductions in negative effects
on soil and the overall environment, respectively [238].
If humanity is to maintaining global feed security in the face of both
rising world population and increasing meat consumption it is essential
to increase crop yields from what is essentially a static or even shrinking
arable land base. The use of GM technologies could allow higher levels
of plant photosynthetic activity, altered plant canopy structure leading
to improved crop light interception, and greater/more efficient alloca­
tion of resources (including mineral nutrients and reduced carbon) from
source to food-related sinks [5,239]; such modifications have the po­
tential to improve the production of biomass yield, either in biomass
crops or as residues from food crops, while allowing the required levels
M. Antar et al.
of global food production. Thus, GM technologies have the capacity to
manipulate appropriate gene expression and allow biomass production
at the levels needed to sustain the developing bioeconomy worldwide
[236].
4.3.2. Genome editing for crop production improvement
Genome editing can accurately and effectively target the genome of
an organism to achieve a desired goal, including improvement of agri­
cultural traits and enhancement of plant resistance [235]. Recently, the
clustered regularly interspaced short palindromic repeats (CRISPR)/Cas
system has become prevalent since it is easier to apply than the earlier
genome editing approaches; it reduces the potential for effects on
untargeted genes [240]. In addition, developing transgenic plants with
the CRISPR-Cas9 system could provide non-transgenic genome modifi­
cations when it hybridizes with current high-performing agricultural
plant species, in the absence of the negative effects perceived to be in
currently commercialized transgenic plants [241]. Moreover, the
CRISPR-Cas9 system can contribute to biomass production improvement
through introduction of elite genes. This has been verified for various
crops including wheat [242], maize [243], sorghum [244] and Brassica
species [245]. Furthermore, the CRISPR-Cas9 system also plays a key
role in the enhancement of resistance to abiotic and biotic stresses,
which decrease quality and yield of crops. For instance, the application
of CRISPR/Cas9 technology helps rice tolerate the damage caused by
powdery mildew, leading to greater yield and quality, through triggered
expression of the TaEDR1 gene [246].
Therefore, manipulating the expression of genes could be a feasible
strategy to enhance crop growth and biomass yield, such as manipu­
lating the expression of gene related nitrogen uptake, assimilation and
transport which can improve the nitrogen use efficiency of plants, which
generally benefits crop yield production and the associated environ­
ment. CRISPR/Cas9, is a promising method to edit targeted genes for
improving plant biomass production; it has already been used in agri­
cultural production [247]. It has been found that overexpression of
transporters in the NRT1 and NRT2 families can enhance yield pro­
duction through regulating the N transport process to improve nitrogen
use efficiency [248]. Similar results have also been reported by Fan et al.
[249]. It has also been found [250] that gene editing also can stimulate
the P uptake to enhance plant development and growth.
The effects of plant biotechnologies are not only on non-food crops,
they also have the ability to improve the production of new crop plant
types [251]. Genome editing has a critical effect in developing new
bio-energy crops, which could contribute optimum biomass production
on marginal land, or even on wastelands, and in developing crops with
resilience to polluted soil and climate change [252].
5. Discussion
There is a critical need to investigate and improve biomass produc­
tion through all available approaches and develop new technologies,
making it possible to feed a growing global population, and provide
associated bioenergy and high value products associated with a bio­
economy approach. The world is changing in concerning ways, with
intense environmental limitations narrowing the global yield capacity of
agricultural systems [253]. In more extreme scenarios, there would be
an ultimate need to find a natural and/or sustainable ways of increasing
crop productivity by assisting plants to withstand extreme environ­
mental uncertainties. To meet global food demand both for humans and
livestock, food production and supply must be doubled by 2050.
Although, global food production is increasing, instead, increased crop
utilization in non-food purposes (first generation biofuels) can interfere
with food supply targets worldwide [254]. There are mixed opinions in
the literature regarding “food vs fuel” indicating how biofuel production
affects global food supply and economy. On one hand, it was suggested
that increased biofuel production reduced food supply by using food
materials as feedstock for renewable fuel generation [255]. On the other
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Renewable and Sustainable Energy Reviews 139 (2021) 110691
hand, biofuel production has been applauded for increased crop pro­
duction and land utilization, and its environmentally friendly aspects
[256]. However, in both cases, the food production should be increased
sufficiently to supply required inputs to both food and biofuel industries.
Under the current situation, adaptations to soil resources will be
likely to provide short-term improvements to increase biomass pro­
duction by using existing technologies. As discussed above, fertilizer
application, especially nitrogen, has positive effects on biomass pro­
duction, but also has negative side effects with regard to environmental
pollution. Thus, application of these approaches to aspects of nitrate
transport and assimilation, combined with better fertilizer application
practices (e.g. precision agriculture) and overall enhanced field man­
agement, it could also be possible to reduce global consumption of ni­
trogen fertilizer while maintaining, or even increasing, crop yield [248].
However, the improvement of biomass production is complex in that it
could be affected by the stress conditions, such as drought/flooding and
heat/cold, even those associated with excessive fertilization input.
Natheless, an important ecosystem component, the rhizosphere is often
ignored. The rhizosphere contains a complex biological community,
comprised of bacteria, fungi, and a wide range of other organisms [257].
The abundant bio-components in the soil have a significant effect on the
growth and development of plants. Most research in this area has been
directed toward the exploration of beneficial microbes from biomass
yield improvement, to disease resistance, to enhancement of overall
resilience to climate change [34,258]. Researchers are considering
alternative techniques that could sustainably increase crop production,
including the utilization of the phytomicrobiome, which is now recog­
nized as having the potential to underpin a “fresh” green revolution
[259]. The application of specific phytomicrobiome members onto food
crops has been studied to a reasonable degree, however, by comparison,
their utilization on energy crops has rarely been considered, despite the
fact that they could be equally useful and efficient in that context [260].
For instance, the inoculation of Azotobacter and Azospirillum increased
the oil quality of safflower (a potential energy crop) by reducing acid
value, iodine value and free fatty acid (oleic acid) contents. In addition,
the microbial inoculants reduced the need for fertilizer inputs (N and P)
by 50–70% [261]. In the same way, Azospirillum has improved the
quality of biodiesel generated from canola seeds, by reducing the acid
value and free fatty acid content. However, the oil obtained from inoc­
ulated and fertilizer supplemented plants were 92 and 93% converted to
methyl esters, respectively [262]. Similarly, biofertilizers (Rhizobium
plus phosphate solubilizing microbes) when compared with chemical
fertilizers, decreased the acid value and free fatty acid content of soy­
bean oil, as compared to the control treatment; plant seed oils supple­
mented with biofertilizers exhibited maximum conversion to methyl
esters [263]. Also, rhizobacterial strains isolated from Brassica species
increased growth and yield, and oil content of canola seeds by 7–57%
[264]. The role of PGPR in stress-amelioration (drought, salinization,
pathogens) are also discussed in a previous research paper, with the
perception being of their importance and efficacy in agricultural sys­
tems, even under more extreme environmental conditions [265].
Application of key phytomicrobiome members to food crops, resulting in
increased yield of food plus increased production of crop residues, a
portion of which could be used in biofuel production, shifts the situation
from one of food versus fuel to food and fuel. In addition, its utilization
in energy-based cropping systems may reduce total production costs by
increasing yield, and reducing fertilizer and pesticide inputs, addressing
one of the major concerns of biofuel production, since 75% of the total
feedstock cost is related to the value of production inputs [266].
Overall enhancement of N assimilation has the capacity to enhance
the yield of key crop species, allowing for improved short- and long-term
nitrogen use efficiency [248]; this could be achieved through modifi­
cation of existing nitrogen-metabolism-related enzymes and the meta­
bolic pathways they play roles in, thus exerting effects at the molecular,
cellular and, finally, whole plant levels. This is based on aspects of crop
gene makeup and regulation; the nutrient uptake and stress
M. Antar et al.
management, leading to more efficient and more effective crop and,
from the perspective of the bioeconomy, biomass production. However,
the benefits provided by associated microbes, including PGPR and
arbuscular mycorrhiza, depend on the attributes and traits of the mi­
crobes, which cause the complex plant-growth-enhancement phenom­
enon, since it not only depends on the microbial species but also the host
plant. Those benefits achieved through microbial- or plant-signal pro­
duction are involved in a large range of plant-microbe interactions
[267]. The best example is the symbiotic interaction between legume
plants and rhizobia. In this case, an isoflavonoid signal secreted from
legume plants stimulates the production of LCO signals from rhizobia,
which can be returned to legume plants and cause commencement of the
nodulation process, leading to the eventual onset of nitrogen fixation in
the symbiosis [268,269]. The appropriate signal released by the correct
microbe, in association with the correct host plants can result in opti­
mum benefits to both of plants and microbes. Thus, one potentially
high-impact approach is to make specialist signals, such as those of the
legume-rhizobia N2-fixing symbiosis, become generalist signals, in order
that they can be applied to a range of plants. Recently, it has been shown
that the LCO rhizobia-to-legume signal has a generalist role in that it can
enhance stress tolerance of a very wide range of legume and non-legume
plants, so that adding the correct isoflavonoid signal to rhizobia can
result in LCO production for generalist application [270]. In terms of
identification of common signals needed for a set range of plants and
isolation of those signals from beneficial microbes, suitable progress in
this area may well help other specialist signals to be commercialized and
deployed in a general context, to improve overall biomass production
[267,271].
Another successful signal molecule produced by members of phyto­
microbiome is thuricin 17; a single small peptide produced from
B. thuringiensis NEB17, a bacterium isolated from soybean root nodules
in our laboratory [272]. This signal molecule acts as a bacteriocin
against other, often closely related, microbial strains, making it a
bacteriocin and not an antibiotic. Thuricin 17 has no antimicrobial ac­
tivity to nodulating rhizobia and plant growth promoting rhizobacteria
[273]. Interestingly, several experiments indicated that thuricin 17
stimulates plant growth and development, leading to more biomass
production, particularly under stressful conditions, through direct and
indirect mechanisms. Inhibitory effects on plant-root pathogens,
resulting in more healthy plants, and increased plant pathogen resis­
tance induction is an indirect mechanism of benefit to plant growth [24,
274]. Having a synergistic collaboration between bacteriocin producers
and nodulating bacteria could be another plant growth promoting
mechanism [274]. When this molecule binds to the receptor in plant
leaves or roots tissues, it acts as pseudo-stress signal that stimulates
specific metabolic pathways, such as increases in photosynthetic rate,
particularly under stressful conditions. In the case of thuricin 17, the
response can be activated without any real stress, causing increases in
net growth enhancement [275]. So far, thuricin 17 has been evaluated
on a range of crops such as corn, soybean and canola and either stim­
ulates plant growth or enhances resistance to abiotic stresses [274,
276–279]. Inoculation with these compounds represents a promising
approach to reduced dependency on fertilizers and/or pesticides, which
could lead to reduced GHG emissions associated with biomass produc­
tion and cause improved total yield.
The interaction between genotype and environment on the expres­
sion of targeted traits should also be considered, since any variation
could be a significant factor affecting genotype productivity under field
conditions. Alternative biomass cropping systems could be another
challenge, since such systems need high bioenergy outputs and low in­
puts. It could be costly to transfer to new cropping systems, such as from
a flexible annual cropping system to a perennial system, which take
more time to achieve profitability. So, to address those challenges,
laboratory work must be extended to field evaluations and eventually
become adopted agricultural practices, since some growth conditions
are largely uncontrollable and are becoming more extreme due to
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climate change. In addition, biomass production, crop adaptation to the
marginal lands and overall more effective cropping systems are needed
to achieve current goals for the bioeconomy. Biofuels and bioproducts
are now considered as “green technologies”, however, details of the
bioenergy crop itself is still largely ignored by crop producers.
In summary, it is crucial to conduct controlled experiments evalu­
ating environmental conditions to understand crop responses to various
stress, which have huge impacts on plant growth and lower overall crop
productivity. These experiments will expand the understanding of the
growth, development and productivity of crop species produced under
field conditions. In addition, attempts should be made, under field
conditions, to select crops that produce the highest biomass and which
ensure production of maximum amount of biomass for food, feed and
biofuel commercialization. More genetic (manipulation and trans­
formation) studies are also required to produce crops tolerant of envi­
ronmental stresses (biotic and abiotic) resulting in greater biomass
accumulation. Also, advancement and deployment of cutting-edge
research technologies is an excellent way to allow in depth investiga­
tion of crop plants at, for instance, the molecular level and to understand
metabolic pathways underpinning biomass accumulation and produc­
tion. These technologies include, but are not limited to, phenomics using
imaging and screening advances, and the various “omics” such as
metabolomics, transcriptomics and proteomics.
6. Conclusions
In coming years, energy production will demand increasing amounts
of biomass. This demand could lead to a gap between biomass produc­
tion and potential consumption, possibly starting as soon as 2020. For
energy production purposes, forest residues, agricultural biomass and
purpose grown biomass crops will play key roles as biomass feedstock.
Many producers and commercial entities are planning to obtain
immediate rewards, especially with regard to increasing of production
inputs. Renewable energy and biomass production are promising areas;
this is a fast-moving area which requires attention and effort, therefore a
review of our current understanding of the area is required, to improve
current overall understanding and build on the knowledge on existing
green technologies and awareness of their potential value. The contri­
bution of this work is provision of an understanding regarding produc­
tion and utilization of biomass around the world. Our work shows that
biomass utilization is receiving more and more attention in most
developed and developing countries, even if some work is at an early
stage, and research is still ongoing regarding the improvement of
biomass production that aims to decrease inputs and increase the
output. In general, novel technologies could face a large variety of
challenges and barriers, but additional research initiatives, collabora­
tions and overall global effort could provide the potential and com­
mercial value required.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgment
The authors would like to acknowledge the support for this review
paper was provided through the Biomass Canada Cluster (BMC), which
is funded through Agriculture and Agri-Food Canada’s AgriScience
program and industry partners.
M. Antar et al. Renewable and Sustainable Energy Reviews 139 (2021) 110691

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